205 research outputs found
Globalization, Universities and Sustainable Human Development
Robert Forrant and Jean L. Pyle look at the fast changing global economy and discuss what can be the role for the university, as an institution, to promote sustainable human development. They contend that the university is an important civil society institution, whose power could be considerable in promoting sustainability. The authors explore ways the university can play a role in fostering a development approach that countervails the preponderant power of institutions advocating a ‘market-driven’ development strategy. Cohesive links at the local and regional level between higher education and industry and community-based organizations create the possibility that difficult societal problems will be tackled and resolved. Development (2002) 45, 102–106. doi:10.1057/palgrave.development.1110388
Chromis degruyi Pyle, Earle & Greene, 2008, new species
Chromis degruyi, new species urn:lsid:zoobank.org:act: 1859 B 68 B- 340 C- 44 F 9 -BEAB-D 75 BAED 300 F 2 DeGruy’s Chromis (Figs. 4 a – 4 c; Table 5; Morphbank 122; DigiMorph 123; GenBank 124; Barcode 125) Holotype. BPBM 40842 126 (81.0 mm SL), Belau (Palau) Islands; Kayangel Atoll, W side; on outer reef dropoff near tip of small reef extension (8 ° 4 ' 16.64 "N, 134 ° 40 ' 54.52 "E): rocky ledge with holes at base of steep sandy slope with many gorgonians, 85 m, hand net, R.L. Pyle, 22 April 2007 [PCMB 3086 127]. Paratypes. BMNH 2007.10. 31.4 128 (38.7 mm SL), Caroline Islands; Yap, S end; “Magic Kingdom” (9 ° 26 ' 3.41 "N, 138 ° 2 ' 5.96 "E): deep rubble on rocky slope, 85 m, quinaldine and hand net, R.L. Pyle, 20 April 2007 [PCMB 3084 129]. CAS 225758 130 (38.3 mm SL), Caroline Islands; Puluwat Atoll; Alet Islet, S side (7 ° 21 ' 15.44 "N, 149 ° 10 ' 47.03 "E): outer reef drop-off with small caves and holes, 100–103 m, quinaldine and hand net, R.L. Pyle and B.D. Greene, 11 April 2007 [PCMB 3032 131]. USNM 391139 132 (76.6 mm SL), Belau (Palau) Islands; off Ngemlis Island; below and slightly to the N of the Blue Holes cave system (7 ° 8 ' 16.49 "N, 134 ° 13 ' 18.5 "E): in coral and rubble at the base of a large boulder offset from the drop-off, 88 m, hand net, R.L. Pyle, 27 April 2007 [PCMB 3114 133]. WAM P. 32901 -001 134 (82.4 mm SL), Belau (Palau) Islands; Ngaruangl Atoll, S end (8 ° 8 ' 50.39 "N, 134 ° 37 ' 3.47 "E), 115 m, hand net, R.L. Pyle, 23 April 2007 [PCMB 3088 135]. Diagnosis. Dorsal rays XIII–XIV, 11–12 (usually XIV, 12); anal rays II, 11–12 (usually 12); pectoral rays 18; spiniform caudal rays 3; tubed lateral-line scales 15–17; gill rakers 7 + 20–21 (total 27–28); body depth 1.84–1.99 in SL; color of adults when fresh dull brownish yellow with nine thin lavender-gray stripes on side of body, with a prominent black spot on dorsal half of pectoral-fin base. Description. Dorsal rays XIV, 12 (one paratype with XIII, another with 11); anal rays II, 12 (one paratype with 11); all dorsal and anal rays branched, the last to base in some specimens; pectoral rays 18, the upper 2 and lowermost unbranched; pelvic rays I, 5; principal caudal rays 8 + 7 = 15; upper and lower procurrent caudal rays 5, the anterior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16 | 15 (15–17, one paratype with 17); posterior midlateral scales with a pore or deep pit 8 | 9 (5–9); scales above dorsal fin to origin of dorsal fin 3; scales below lateral line to origin of anal fin 9 (one paratype with 8); gill rakers 7 + 20 = 27 (7 + 20–21 = 27–28); surpaneural (predorsal) bones 3; vertebrae 12 + 13. Body moderately deep, depth 1.84 (1.92–1.99) in SL, and compressed, the width 2.87 (2.73–3.29) in body depth; head length 3.10 (2.95–3.18) in SL; dorsal profile of head with slight convexity anterior to eye, very slight concavity dorsal to eye, and very slight convexity on nape; snout shorter than orbit diameter, its length 4.05 (3.63–4.38) in head length; orbit diameter 2.77 (2.12–2.95) in head length; interorbital space convex, its width 2.73 (2.73–3.15) in head length; caudal-peduncle depth 2.18 (2.11–2.27) in head; caudal-peduncle length 2.83 (2.69–3.37) in head. TABLE 5. Proportional measurements (%SL) and counts of Chromis degruyi, new species. Values separated by a pipe “|” are left|right or upper|lower. Holotype Paratypes Mouth terminal, small, oblique, the upper jaw forming an angle of about 37 º to horizontal axis of head and body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw length 2.91 (3.05–3.18) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly; about 20 upper and about 20 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with rounded tip; gill rakers long and slender, the longest on lower limb near angle about three-fourths length of longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of pupil, slightly less than one-third distance from front of snout to base of upper lip. Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with free lower margin extending nearly to a vertical at posterior edge of pupil. Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 13 th and 14 th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns, progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic spine about one-half length of spine. Origin of dorsal fin over third lateral-line scale, the pre-dorsal distance 2.39 (2.30–2.41) in SL; base of spinous portion of dorsal fin contained 2.17 (2.14–2.34) in SL; base of soft portion of dorsal fin contained 6.66 (6.54–7.17) in SL; first dorsal spine 12.33 (9.51–12.14) in SL; second dorsal spine 8.15 (6.84–7.96) in SL; third dorsal spine 5.88 (5.53–6.39) in SL; fourth dorsal spine 5.23 (5.04–5.87) in SL; fifth dorsal spine 5.08 (5.17–5.76) in SL; sixth dorsal spine 5.07 (4.93–5.79) in SL; last dorsal spine 6.27 (6.14–7.04) in SL; membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.95 (4.54–5.04) in SL; first anal spine 11.30 (10.99–13.08) in SL; second anal spine 3.84 (3.76–4.52) in SL; first anal soft ray the longest, its length 4.48 (4.34–5.15) in SL; caudal fin forked, its length 2.84 (2.26–3.29) in SL, the caudal concavity 5.63 (4.37–5.46) in SL; fourth pectoral-fin ray longest, 2.99 (2.79–3.20) in SL; pelvic spine 5.36 (5.66–6.12) in SL; first soft ray of pelvic fin filamentous, usually reaching to first or second analfin ray (when not broken or otherwise damaged), its length 2.94 (2.95–4.17) in SL. Color of adults when fresh dull brownish yellow with nine thin lavender-gray stripes, some faint, the middle 4 or 5 extending onto caudal peduncle; nape area olive-brown, lighter on thorax and ventrally to anus, becoming yellowish white; black spot slightly smaller than orbit at upper pectoral axil; faint diffuse lavender blotch smaller than orbit on opercle edge at level of lower orbit, not apparent underwater; olivaceous with brown stripes and greenish olive in nape area when observed underwater; spinous portion of dorsal fin olivebrown, distal one-fourth yellowish white; basal half of soft dorsal fin dark brown with almost black outer margin, distal half very light yellowish white to translucent on some specimens; caudal fin olive-brown, inner rays yellowish white; anal fin spines yellowish white, rays and membranes on basal half light olive-brown becoming distally yellowish white; black blotch smaller than orbit centered in posterior distal anal fin, more apparent in large specimens; pectoral fin translucent; pelvic-fin spine and first ray white, successive rays and membranes on basal half olive-brown, distal half yellowish white; iris brownish yellow; juveniles bluish gray; a bright yellow blotch on the distal half of the soft dorsal fin, covering the second through fifth dorsal soft rays, rays 6 to last paler than anterior part of soft dorsal fin; a bright yellow stripe from posterior base of soft dorsal fin in a band approximately the width of 2 scales continuing dorsally to tip of outer rays of dorsal lobe of caudal fin; lower caudal rays with a similar yellow band commencing ventrally on caudal peduncle and extending to distal tip of lower caudal fin rays, approximately 7 median caudal rays white; third through seventh anal-fin rays and intervening membranes bright yellow on distal third, succeeding rays white. Color in alcohol similar to fresh color, except paler brown overall. Distribution. Observed or collected throughout the Caroline Islands, from Puluwat to Palau. A Chromis resembling the juvenile of this species was observed in Fagatele Bay in May 2001 by the senior author. Etymology. Named degruyi to honor Michael V. DeGruy, in recognition of the sincere enthusiasm and determination he demonstrated while attempting to collect the first adult specimen of this species. Remarks. The habitat of this species is similar to that of other species described herein: deep outer-reef slopes at depths of 85–120 m, usually in the vicinity of rock outcrops with small holes and caves, and around limestone talus. It is generally not as abundant as other species of Chromis described here, usually found in small groups and observed feeding low in the water column. The species appears most similar to other deep-dwelling species of Chromis described previously (see Remarks section of C. abyssus). Juveniles superficially resemble C. opercularis 136 (Günther in Playfair and Günther 1867) in color, but are easily distinguished from that species on several morphological characters (e.g., usually XIV dorsal-fin spines in C. degruyi, vs. XIII in C. opercularis; body depth 1.84–1.99 in SL vs. 2.1–2.3 in C. opercularis), as well as adult coloration. Some aspects of the adult coloration are similar to C. planesi Lecchini and Williams 2004, but C. degruyi differs from that species in many other aspects of adult coloration as well as number of pectoral-fin rays (20 in C. planesi vs. 18 in C. degruyi), dorsal-fin soft rays (usually 13 vs. usually 12), and tubed lateral-line scales (17 vs. usually 15–16).Published as part of Richard L. Pyle, John L. Earle & Brian D. Greene, 2008, Five new species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the tropical western Pacific, pp. 3-31 in Zootaxa 1671 on pages 18-21, DOI: 10.5281/zenodo.18018
Chromis abyssus Pyle, Earle & Greene, 2008, new species
Chromis abyssus, new species urn:lsid:zoobank.org:act: 8 BDC0735-FEA 4-4298 - 83 FA-D04F 67 C 3 FBEC Deep Blue Chromis (Figs. 1 a – 1 c; Table 1; Morphbank 59; DigiMorph 60; GenBank 61; Barcode 62) Holotype. BPBM 40861 63 (81.6 mm SL), Belau (Palau) Islands; off Ngemelis Island; below and slightly N of Blue Holes caverns (7 ° 8 ' 16.49 "N, 134 ° 13 ' 18.5 "E): above large rock outcrop, 110 m, hand net, R.L. Pyle, 27 April 2007 [PCMB 3113 64]. Paratypes. BMNH 2007.10. 31.1 65 (50.2 mm SL) [PCMB 3103 66]. BPBM 40855 67 (3; 37.6 –98.0 mm SL) [PCMB 3100 68, 3102 69]. CAS 225755 70 (64.1 mm SL) [PCMB 3105 71]. MNHN 2007.1922 72 (63.7 mm SL) [PCMB 3104 73]. USNM 391136 74 (2; 44.4–90.2 mm SL) [PCMB 3101 75]. WAM P. 32898 -001 76 (64.5 mm SL) [PCMB 3106 77]. All from same locality as holotype: sand and rubble slope with scattered rock outcroppings, 107–116 m, quinaldine and hand net, R.L. Pyle and B.D. Greene, 25 April 2007. Diagnosis. Dorsal rays XIV, 12–13 (usually 13); anal rays II, 12–14 (usually 13); pectoral rays 18–19 (usually 19); spiniform caudal rays 3; tubed lateral-line scales 14–16; gill rakers 6–7 + 17–18 (usually 7 + 18; total 24–25, usually 25); body depth 1.58–1.83 in SL; color when fresh dark gray with a large iridescent dark blue spot at center of each scale; membranes on median fins and pelvic fins opaque charcoal gray, with an iridescent dark blue margin on the spinous portion of the dorsal and anal fins; caudal fin mottled iridescent dark blue and black; pectoral fins with a black ovoid spot covering the basal portion and pectoral-fin axil. Description. Dorsal rays XIV, 13 (two paratypes with XIV, 12); anal rays II, 13 (II, 12–14); all dorsal and anal rays branched, the last to base in some specimens; pectoral rays 19 (one paratype with 18), the upper 2 and lowermost unbranched; pelvic rays I, 5; principal caudal rays 8 + 7 = 15; upper and lower procurrent caudal rays 5, the anterior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16 | 14 (14–16 except for one paratype with 11 | 15); posterior midlateral scales with a pore or deep pit 7 | 6 (5–8); scales above dorsal fin to origin of dorsal fin 3.5 (3–3.5); scales below lateral line to origin of anal fin 9 (9–10); gill rakers 7 + 18 = 25 (6–7 + 17–18 = 24–25), 6 gill rakers on upper limb of gill arch in one paratype, 17 gill rakers on lower limb of gill arch in one paratype); surpaneural (predorsal) bones 3; vertebrae 12 + 13. Body moderately deep, depth 1.73 (1.58–1.83) in SL, and compressed, the width 3.23 (2.65–3.56) in body depth; head length 3.01 (2.79–3.05) in SL; dorsal profile of head with slight convexity anterior to eye, slight concavity dorsal to eye, and slight convexity on nape; snout shorter than orbit diameter, its length 4.11 (4.00– 5.74) in head length; orbit diameter 2.66 (2.28–2.90) in head length; interorbital space convex, its width 2.79 (2.59–2.92) in head length; caudal-peduncle depth 2.17 (2.15–2.59) in head; caudal-peduncle length 3.01 (3.34–4.39) in head. Mouth terminal, small, oblique, the upper jaw forming an angle of about 40 º to horizontal axis of head and body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw length 3.57 (3.39–4.11) in head; an outer row of conical teeth in each jaw, largest anteriorly; about 27 upper and about 20 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with rounded tip; gill rakers long and slender, the longest on lower limb near angle about four-fifths length of longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of pupil, slightly less than one-third distance from front of snout to base of upper lip. Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with free lower margin extending nearly to a vertical at posterior edge of pupil. Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 11 th and 12 th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns, progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic spine about one-half length of spine. Origin of dorsal fin over second lateral-line scale, the pre-dorsal length 2.29 (2.24–2.54) in SL; base of spinous portion of dorsal fin contained 2.24 (2.02–2.39) in SL; base of soft portion of dorsal fin contained 5.75 (5.65–6.48) in SL; first dorsal spine 10.85 (7.78–11.01) in SL; second dorsal spine 6.64 (5.22–7.14) in SL; third dorsal spine 5.41 (4.53–5.42) in SL; fourth dorsal spine 5.00 (4.44–5.08) in SL; fifth dorsal spine 4.88 (4.39–5.05) in SL; sixth dorsal spine 4.90 (4.45–4.99) in SL; last dorsal spine 6.19 (6.11–7.40) in SL; membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, sometimes with a filamentous extension, its length 4.39 (4.21–5.06) in SL; first anal spine 10.92 (8.76–11.13) in SL; second anal spine 4.03 (3.66–4.24) in SL; first anal soft ray the longest, its length 4.41 (4.15–4.62) in SL; caudal fin forked, without significant filamentous extensions, its length 2.89 (2.87–3.64) in SL, the caudal concavity 5.83 (4.54 –8.00) in SL; fourth pectoral-fin ray longest, 2.77 (2.65–3.08) in SL; pelvic spine 5.20 (4.95–5.67) in SL; first soft ray of pelvic fin filamentous, usually reaching to first through third anal-fin ray (when not broken or damaged), its length 2.86 (2.54–4.03) in SL. Color of adults and juveniles when fresh predominantly charcoal gray, a large iridescent dark blue spot at center of each scale (including scales on head and median fins), blue spots occupying about half of visible area of each scale on body, decreasing in size slightly towards abdomen and ventral portion of body, blue spots forming a near-continuous line along base of dorsal and anal fins, a vertical column of scales with iridescent dark blue spots extending dorsally on each interspinous membrane of dorsal fin, blue spots on scales covering soft portions of dorsal and anal fins varying in size, forming a mottled pattern of blue and black; membranes on median fins and pelvic fins opaque charcoal gray, with an iridescent dark blue margin on spinous portion of the dorsal fin, and a broad iridescent dark blue margin on the anal fin; caudal fin mottled iridescent dark blue and black; pelvic-fin spine entirely iridescent dark blue, an iridescent dark blue streak on the pelvic-fin soft rays, the filamentous extension on the pelvic fin white; pectoral fins translucent charcoal gray with a ovoid black spot on base and axil; iris charcoal gray to black; iridescent blue fleshy orbit margin. Color in alcohol similar to general color pattern when fresh, except charcoal gray pigment sometimes fades to brownish gray, and iridescent dark blue is either faded to pale gray blue, or has disappeared altogether (leaving the specimen uniform brownish gray, or sometimes charcoal gray overall). Distribution. Only collected from the type locality; also observed at similar depths at Augulpelu Reef in Palau. An individual of what appears to be this species was observed and photographed by Mr. Forrest Young at 120–150 m near Manado, Sulawesi, Indonesia. Etymology. Named abyssus, a Latinized form of the Greek noun abyssos (meaning “abyss”), to honor the documentary film Pacific Abyss, produced by the British Broadcasting Corporation (BBC), which funded the expedition on which the type specimens were collected. The vernacular name “Deep Blue Chromis ”, a reference to both the life color of this species and the relatively (within the context of the genus) deep-dwelling habits, is suggested instead of the more literally translated “Abyss Chromis ”, so as not to imply that the species inhabits depths commonly defined as “abyssal”. Remarks. This species was first observed by the senior author on May 10, 1997, during a mixed-gas rebreather dive to 120 m on the east side of Augulpelu Reef; Palau (07º 16.41 ' N, 134 º 31.44 ' E). It was later observed at the same reef at depths of 117–139 m from a submersible by Patrick L. Colin and Lori J. Bell in February–March, 2001. In April 2005, Mr. Forrest Young and colleagues observed several individuals of this (or a very similar) species during mixed-gas rebreather dives at depths of 120–150 m at Manado, Sulawesi, Indonesia. The type specimens included herein are the first of this species to be collected. From these observations, C. abyssus appears to prefer depths in excess of 115 m, staying close to the substratum among boulders and rock outcroppings, where it takes refuge in small caves and holes. Juveniles and some subadults were also observed around limestone talus. Adults were usually observed singly or in pairs, while subadults and juveniles were seen in small groups. All type specimens were collected in the same general area, where the species is not uncommon. Other Chromis observed in the vicinity include three of the new species described herein (C. brevirostris, C. degruyi, and C. earina). Chromis abyssus is not obviously allied with any other known species of the genus. It shares some similarities with a group of seven Indo-Pacific deep-dwelling Chromis species, characterized by a similar stout body shape, a large eye, and usually XIV dorsal spines. In their 1985 description of C. abyssicola, Allen and Randall noted a complex of deep-dwelling Chromis species distinguished by, among other characters, 19 or 20 pectoral rays, and 28–34 gill rakers. In addition to C. abyssicola, their complex included C. megalopsis 78 Allen 1976 (now regarded as a junior synonym of C. mirationis Tanaka 1917), C. mirationis Tanaka 1917 and C. struhsakeri Randall and Swerdloff 1973, to which we would add the subsequently named C. planesi 79 Lecchini and Williams 2004. C. abyssus has fewer pectoral rays (18 or 19) and fewer gill rakers (24–28) than members of this species complex, and may comprise a second grouping of deep-dwelling Chromis species, along with C. okamuri Yamakawa and Randall 1989 from Japan, the East African C. woodsi 80 Bruner and Arnam 1979 (both easily distinguished from C. abyssus on the basis of color and certain morphological characters such as number of gill rakers and tubed lateral-line scales), as well as the two new species C. circumaurea and C. degruyi, both described herein. Of the remaining two deep-dwelling Indo-Pacific stout-bodied Chromis species with XIV spines, C. onumai Senou and Kudo 2007 has the high pectoral-ray count of the first complex (19–20) and the gill-raker count of the second (25–27). C. axillaris 81 (Bennett 1831) has a wide gillraker range (26–30) and cannot easily be placed in either complex by this character. Of the three new Chromis with XIV dorsal-fin spines described herein (C. abyssus, C. circumaurea, and C. degruyi), each has a unique and distinctive color pattern, and is readily distinguished from the others. Among the three, the former two (C. abyssus and C. circumaurea) share the most similarities both in terms of morphology and in Barcode DNA sequence data.Published as part of Richard L. Pyle, John L. Earle & Brian D. Greene, 2008, Five new species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the tropical western Pacific, pp. 3-31 in Zootaxa 1671 on pages 6-9, DOI: 10.5281/zenodo.18018
Chromis circumaurea Pyle, Earle & Greene, 2008, new species
Chromis circumaurea, new species urn:lsid:zoobank.org:act:8ADC4817-8F1C-4C88-8B8A-5372A84CAEC9 Gold-rim Chromis (Figs. 3a -3c, Ta b l e 4; Morphbank105; GenBank106; Barcode107) Holotype. BPBM 40836108 (98.2 mm SL), Caroline Islands; Yap, S end; "Magic Kingdom" (9°26'3.41"N, 138°2'5.96"E): among boulders on sloping shelf above deep drop-off, 98-100 m, hand net, R.L. Pyle and B.D. Greene, 20 April 2007 [PCMB 3080109]. Paratypes. BMNH 2007.10.31.3110 (102.4 mm SL) [PCMB 3081111]. CAS 225757112 (97.6 mm SL) [PCMB 3078113]. MNHN 2007-1924114 (92.5 mm SL) [PCMB 3076115]. USNM 391138116 (94.2 mm SL) [PCMB 3077117]. WAM P.32900-001118 (96.6 mm SL) [PCMB 3079119]. All with same data as holotype. Diagnosis. Dorsal rays XIV,12-13 (usually 13); anal rays II, 13-14 (usually 13); pectoral rays 18-19; spiniform caudal rays 3; tubed lateral-line scales 16-17; gill rakers 6-7+20-21 (total 26-27); body depth 1.68-1.86 in SL; color when fresh mahogany brown with bright yellow distally on spinous portion of dorsal fin; soft portion of dorsal fin, caudal fin, and anal fin bright yellow. Description. Dorsal rays XIV,13 (12 in one paratype); anal rays II,13 (14 in one paratype); all dorsal and anal rays branched, the last to base in some specimens; pectoral rays 19 (18-19), the upper 2 and lowermost unbranched; pelvic rays I,5; principal caudal rays 8+7=15; upper and lower procurrent caudal rays 5, the anterior3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16|17 (16-17); posterior midlateral scales with a pore or deep pit 8 (5-8); scales above dorsal fin to origin of dorsal fin 3.5 (3-3.5); scales below lateral line to origin of anal fin 10 (9-10.5); gill rakers 6+21=27 (6-7+20-21= 26-27); surpaneural(predorsal) bones 3; vertebrae 12+13. Body moderately deep, depth 1.71 (1.68-1.86) in SL, and compressed, the width 3.02 (2.93-3.22) in body depth; head length 3.28 (3.18-3.37) in SL; dorsal profile of head with slight convexity anterior to eye, slight concavity dorsal to eye, and slight convexity on nape; snout shorter than orbit diameter, its length 3.71 (3.82-4.30) in head length; orbit diameter 2.61 (2.35-2.63) in head length; interorbital space convex, its width 2.61 (2.54-2.74) in head length; caudal-peduncle depth 2.04 (2.02-2.04) in head; caudal-peduncle length 3.19 (2.72-3.40) in head. Mouth terminal, small, oblique, the upper jaw forming an angle of about 40º to horizontal axis of head and body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw length 3.09 (2.86-3.22) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly; about 32 upper and about 26 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer row, in 2-3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with rounded tip; gill rakers long and slender, the longest on lower limb near angle about three-fourths length of longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of pupil, slightly less than one-third distance from front of snout to base of upper lip. Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with free lower margin extending nearly to a vertical at posterior edge of pupil. Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 12th and 13th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns, progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic spine slightly more than one-third length of spine. Origin of dorsal fin over fourth lateral-line scale, the pre-dorsal distance 2.31 (2.30-2.47) in SL; base of spinous portion of dorsal fin contained 2.14 (2.09-2.18) in SL; base of soft portion of dorsal fin contained 6.16 (5.68-6.35) in SL; first dorsal spine 12.62 (9.91-10.88) in SL; second dorsal spine 6.83 (6.13-6.88) in SL; third dorsal spine 5.52 (4.81-5.53) in SL; fourth dorsal spine 4.88 (4.73-5.09) in SL; fifth dorsal spine 4.86 (4.66-5.02) in SL; sixth dorsal spine 4.82 (4.63-5.08) in SL; last dorsal spine 6.40 (6.13-6.57) in SL; membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.20 (4.18-4.67) in SL; first anal spine 11.98 (10.50-11.05) in SL; second anal spine 3.77 (3.89-4.15) in SL; first anal soft ray the longest, its length 4.18 (4.31-4.56) in SL; caudal fin forked, its length 3.17 (2.68-3.39) in SL, the caudal concavity 5.46 (5.20-6.44) in SL; fourth pectoral-fin ray longest, 2.77 (2.62-2.83) in SL; pelvic spine 5.78 (5.09-5.52) in SL; first soft ray of pelvic fin without long filamentous extension, usually not reaching anal fin, its length 3.89 (3.82-4.20) in SL. Color when fresh mahogany brown, appearing slate brown underwater; lateral line faintly brownish cream-colored; scales below lateral line with faint brownish cream-colored broad center area, forming approximately eight horizontal stripes visible underwater; spinous portion of dorsal fin same color as body, becoming bright yellow distally on first spine; second through last dorsal spines and membranes abruptly yellow distally, yellow portion increasing from distal one-fourth of fin at third spine to distal half at eleventh spine; soft dorsal fin entirely bright yellow except for posteriorly diminishing thin brown area basally on anterior6 rays; caudal region from posterior base of dorsal fin to posterior tip of caudal fin uniform bright yellow; brown body color extends posterior to anal fin to lower anterior caudal peduncle; anal fin spines yellowish white; anal fin rays and membranes bright yellow; scales along ventral margin from anus to origin anal fin yellow; pectoral fin translucent; pelvic-fin spine translucent, medial yellow wash on anterior 3 pelvic rays, rays otherwise translucent; pelvic-fin membranes mahogany brown basally, translucent distally; iris brown with yellow wash. Color in alcohol similar to fresh color, except yellow portions are much paler yellow, and brown portions are slightly paler brown (much paler brown on thorax). Distribution. Observed from submersibles in the Marshall Islands and Mariana Islands, but only collected from Yap. Etymology. Named circumaurea, an adjective derived from the Latin words circum (meaning "around") and aurea (meaning "golden, of gold"), in reference to the golden-yellow anal fin, caudal fin, and outer margin of the dorsal fin. Remarks. This species was first observed and photographed from a submersible by Patrick L. Colin at Enewetak in the Marshall Islands. An unconfirmed sighting and video clip of this species from the Mariana Islands requires verification. It was observed at Yap at depths of 98-120 m, in a group of about a dozen individuals living among large (~1-2 m) rock boulders just above the upper edge of a precipitous drop-off. A juvenile of approximately 40 mm SL was observed by the first author at a depth of 120 m, below the site where the type specimens were collected; its color pattern was consistent with that of the adults. Two photos appearing on p. 390 of Kuiter & Tonozuka (2001), labelled as Chromis analis 120 (Cuvier 1830), bear a remarkable resemblance to C. circumaurea, but differ in number of dorsal-fin spines (XIII vs. XIV) and color of body (paler in C. analis), caudal peduncle (dark centrally vs. entriely yellow), and central region of caudal fin (transparent vs. yellow). This species also bears a superficial resemblance in color to Chromis flavicauda121 (Guenther 1880) from the western Atlantic Ocean, but is readily distinguished from that species on the basis of body color (blue in C. flavicauda vs. brown in C. circumaurea), dorsal-fin rays (XIII,11-12 vs. XIV,12-13, usually 13), anal-fin soft rays (11 vs. 13). Similarities with other deep-dwelling species with XIV dorsal-fin spines, including the new species C. abyssus described herein, are discussed in the Remarks section of C. abyssus.Published as part of Pyle, R. L., Earle, J. L. & Greene, B. D., 2008, Five new species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the tropical western Pacific., pp. 3-31 in Zootaxa 1671 on pages 15-1
Chromis circumaurea Pyle, Earle & Greene, 2008, new species
Chromis circumaurea, new species urn:lsid:zoobank.org:act: 8 ADC 4817 - 8 F 1 C- 4 C 88 - 8 B 8 A- 5372 A 84 CAEC 9 Gold-rim Chromis (Figs. 3 a – 3 c, Table 4; Morphbank 105; GenBank 106; Barcode 107) Holotype. BPBM 40836 108 (98.2 mm SL), Caroline Islands; Yap, S end; “Magic Kingdom” (9 ° 26 ' 3.41 "N, 138 ° 2 ' 5.96 "E): among boulders on sloping shelf above deep drop-off, 98–100 m, hand net, R.L. Pyle and B.D. Greene, 20 April 2007 [PCMB 3080 109]. Paratypes. BMNH 2007.10. 31.3 110 (102.4 mm SL) [PCMB 3081 111]. CAS 225757 112 (97.6 mm SL) [PCMB 3078 113]. MNHN 2007 - 1924 114 (92.5 mm SL) [PCMB 3076 115]. USNM 391138 116 (94.2 mm SL) [PCMB 3077 117]. WAM P. 32900 -001 118 (96.6 mm SL) [PCMB 3079 119]. All with same data as holotype. Diagnosis. Dorsal rays XIV, 12–13 (usually 13); anal rays II, 13–14 (usually 13); pectoral rays 18–19; spiniform caudal rays 3; tubed lateral-line scales 16–17; gill rakers 6–7 + 20–21 (total 26–27); body depth 1.68–1.86 in SL; color when fresh mahogany brown with bright yellow distally on spinous portion of dorsal fin; soft portion of dorsal fin, caudal fin, and anal fin bright yellow. Description. Dorsal rays XIV, 13 (12 in one paratype); anal rays II, 13 (14 in one paratype); all dorsal and anal rays branched, the last to base in some specimens; pectoral rays 19 (18–19), the upper 2 and lowermost unbranched; pelvic rays I, 5; principal caudal rays 8 + 7 = 15; upper and lower procurrent caudal rays 5, the anterior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16 | 17 (16–17); posterior midlateral scales with a pore or deep pit 8 (5–8); scales above dorsal fin to origin of dorsal fin 3.5 (3–3.5); scales below lateral line to origin of anal fin 10 (9–10.5); gill rakers 6 + 21 = 27 (6–7 + 20–21 = 26–27); surpaneural (predorsal) bones 3; vertebrae 12 + 13. Body moderately deep, depth 1.71 (1.68–1.86) in SL, and compressed, the width 3.02 (2.93–3.22) in body depth; head length 3.28 (3.18–3.37) in SL; dorsal profile of head with slight convexity anterior to eye, slight concavity dorsal to eye, and slight convexity on nape; snout shorter than orbit diameter, its length 3.71 (3.82–4.30) in head length; orbit diameter 2.61 (2.35–2.63) in head length; interorbital space convex, its width 2.61 (2.54–2.74) in head length; caudal-peduncle depth 2.04 (2.02–2.04) in head; caudal-peduncle length 3.19 (2.72–3.40) in head. Mouth terminal, small, oblique, the upper jaw forming an angle of about 40 º to horizontal axis of head and body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw length 3.09 (2.86–3.22) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly; about 32 upper and about 26 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with rounded tip; gill rakers long and slender, the longest on lower limb near angle about three-fourths length of longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of pupil, slightly less than one-third distance from front of snout to base of upper lip. TABLE 4. Proportional measurements (%SL) and counts of Chromis circumaurea, new species. Values separated by a pipe “|” are left|right or upper|lower. Holotype Paratypes Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with free lower margin extending nearly to a vertical at posterior edge of pupil. Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 12 th and 13 th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns, progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic spine slightly more than one-third length of spine. Origin of dorsal fin over fourth lateral-line scale, the pre-dorsal distance 2.31 (2.30–2.47) in SL; base of spinous portion of dorsal fin contained 2.14 (2.09–2.18) in SL; base of soft portion of dorsal fin contained 6.16 (5.68–6.35) in SL; first dorsal spine 12.62 (9.91–10.88) in SL; second dorsal spine 6.83 (6.13–6.88) in SL; third dorsal spine 5.52 (4.81–5.53) in SL; fourth dorsal spine 4.88 (4.73–5.09) in SL; fifth dorsal spine 4.86 (4.66–5.02) in SL; sixth dorsal spine 4.82 (4.63–5.08) in SL; last dorsal spine 6.40 (6.13–6.57) in SL; membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.20 (4.18–4.67) in SL; first anal spine 11.98 (10.50–11.05) in SL; second anal spine 3.77 (3.89–4.15) in SL; first anal soft ray the longest, its length 4.18 (4.31–4.56) in SL; caudal fin forked, its length 3.17 (2.68–3.39) in SL, the caudal concavity 5.46 (5.20–6.44) in SL; fourth pectoral-fin ray longest, 2.77 (2.62–2.83) in SL; pelvic spine 5.78 (5.09–5.52) in SL; first soft ray of pelvic fin without long filamentous extension, usually not reaching anal fin, its length 3.89 (3.82–4.20) in SL. Color when fresh mahogany brown, appearing slate brown underwater; lateral line faintly brownish cream-colored; scales below lateral line with faint brownish cream-colored broad center area, forming approximately eight horizontal stripes visible underwater; spinous portion of dorsal fin same color as body, becoming bright yellow distally on first spine; second through last dorsal spines and membranes abruptly yellow distally, yellow portion increasing from distal one-fourth of fin at third spine to distal half at eleventh spine; soft dorsal fin entirely bright yellow except for posteriorly diminishing thin brown area basally on anterior 6 rays; caudal region from posterior base of dorsal fin to posterior tip of caudal fin uniform bright yellow; brown body color extends posterior to anal fin to lower anterior caudal peduncle; anal fin spines yellowish white; anal fin rays and membranes bright yellow; scales along ventral margin from anus to origin anal fin yellow; pectoral fin translucent; pelvic-fin spine translucent, medial yellow wash on anterior 3 pelvic rays, rays otherwise translucent; pelvic-fin membranes mahogany brown basally, translucent distally; iris brown with yellow wash. Color in alcohol similar to fresh color, except yellow portions are much paler yellow, and brown portions are slightly paler brown (much paler brown on thorax). Distribution. Observed from submersibles in the Marshall Islands and Mariana Islands, but only collected from Yap. Etymology. Named circumaurea, an adjective derived from the Latin words circum (meaning “around”) and aurea (meaning “golden, of gold”), in reference to the golden-yellow anal fin, caudal fin, and outer margin of the dorsal fin. Remarks. This species was first observed and photographed from a submersible by Patrick L. Colin at Enewetak in the Marshall Islands. An unconfirmed sighting and video clip of this species from the Mariana Islands requires verification. It was observed at Yap at depths of 98–120 m, in a group of about a dozen individuals living among large (~ 1–2 m) rock boulders just above the upper edge of a precipitous drop-off. A juvenile of approximately 40 mm SL was observed by the first author at a depth of 120 m, below the site where the type specimens were collected; its color pattern was consistent with that of the adults. Two photos appearing on p. 390 of Kuiter & Tonozuka (2001), labelled as Chromis analis 120 (Cuvier 1830), bear a remarkable resemblance to C. circumaurea, but differ in number of dorsal-fin spines (XIII vs. XIV) and color of body (paler in C. analis), caudal peduncle (dark centrally vs. entriely yellow), and central region of caudal fin (transparent vs. yellow). This species also bears a superficial resemblance in color to Chromis flavicauda 121 (Günther 1880) from the western Atlantic Ocean, but is readily distinguished from that species on the basis of body color (blue in C. flavicauda vs. brown in C. circumaurea), dorsal-fin rays (XIII, 11–12 vs. XIV, 12–13, usually 13), anal-fin soft rays (11 vs. 13). Similarities with other deep-dwelling species with XIV dorsal-fin spines, including the new species C. abyssus described herein, are discussed in the Remarks section of C. abyssus.Published as part of Richard L. Pyle, John L. Earle & Brian D. Greene, 2008, Five new species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the tropical western Pacific, pp. 3-31 in Zootaxa 1671 on pages 15-18, DOI: 10.5281/zenodo.18018
Scientific Management as Applied to the Personnel Department
It is the intention of the author to offer in this thesis a new point of view in dealing wit~ men in industry. Modern executives are coming ·to realize that their workers are not a ·bulk mass but a group of individuals. They are recognizing, that these workers differ in things they are fitted to do and are capable of doing; also that they differ in their interests, ambitions, and the thing that seem most desirable; and as men of differing capacities and desires they require individual adjustment to the opportunities offered them in the field of industry
Biological Technical Publication BTP-R6011-2008
This is a description of the molt and aging criteria for four grassland passerines. The species covered are Sprague's Pipit, Grasshopper sparrow, Baird's sparrows, and Chestnut-collared Longspurs. Molt information is important for determining demographic parameters, which is key for helping determine causes of population decline in passerines.U.S. Fish & Wildlife Service
Molt and Aging Criteria
for Four North American
Grassland Passerines
Biological Technical Publication
BTP-R6011-2008
Bob Gress©
U.S. Fish & Wildlife Service
Molt and Aging Criteria
for Four North American
Grassland Passerines
Biological Technical Publication
BTP-R6011-2008
Peter Pyle1
Stephanie L. Jones2
Janet M. Ruth3
1 The Institute for Bird Populations, Point Reyes Station, CA
2 U.S. Fish and Wildlife Service, Nongame Migratory Bird Program Region 6,
Denver, CO
3 U.S. Geological Survey, Arid Lands Field Station, Albuquerque, NM
Cover image: Sprague’s Pipit
Photo credit: Bob Gress©
ii Molt and Aging Criteria for Four North American Grassland Passerines
Author contact information:
Peter Pyle
The Institute for Bird Populations
P.O. Box 1346
Point Reyes Station, CA 94956
Phone: (415) 663-2053
E-mail: [email protected]
Stephanie L. Jones
U.S. Fish and Wildlife Service
Region 6 Nongame Migratory Bird Coordinator
P.O. Box 25486 DFC
Denver, CO 80225
Phone: (303) 236-4409
E-mail: [email protected]
Janet M. Ruth
Research Ecologist
U.S. Geological Survey
Fort Collins Science Center
Arid Lands Field Station
Biology Department MSC03 2020
1 University of New Mexico
Albuquerque, NM 87131-0001
Phone: (505) 346-2870
E-mail: [email protected]
For additional copies or information, contact:
U.S. Fish and Wildlife Service
Region 6 Nongame Migratory Bird Coordinator
P.O. Box 25486 DFC
Denver, CO 80225
Recommended citation:
Pyle, P., S.L. Jones, and J. M. Ruth. 2008. Molt and
aging criteria for four North American grassland
passerines. U.S. Department of Interior, Fish and
Wildlife Service, Biological Technical Publication,
FWS/BTP-R6011-2008, Washington, D.C.
Table of Contents iii
Table of Contents
List of Figures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv
Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Molt and Aging by Molt Limits. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Species Accounts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Sprague’s Pipit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Grasshopper Sparrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Baird’s Sparrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Chestnut-collared Longspur. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Appendix. Feather Collection Protocols for Analysis (Isotope and DNA). . . . . . . . . . . . . . . . . . . . . . 19
iv Molt and Aging Criteria for Four North American Grassland Passerines
List of Figures
Figure 1. . Molt patterns among four species of grassland passerines during the first and definitive molt
cycles. Solid bars indicate complete molts and dashed bars indicate incomplete, limited, or partial molts.
Locations of where molts occur are indicated above the bars: BG = breeding grounds; WG = wintering
grounds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Figure 2. Tracts of the wing for identification of molt limits in grassland passerines. The primary covert tract
is emphasized because it is often the best tract to use to separate SYs from ASYs in spring. Primaries are
numbered distally and secondaries are numbered proximally, reflecting the order in which they molt (except
for the tertials, which molt before other secondaries). Note that the “tertials” are usually considered a subset
of the secondaries, s7-s9. Illustration: Steve N.G. Howell . .2
Figure 3. Indication of molt limits within the wings of grassland passerines. Darker feathers indicate those
replaced during the preformative (or prealternate) molt. A-E indicate limits resulting from varying extents
of these molts. F represents uniform feather tracts following a complete molt. Illustration: Steve N.G.
Howell. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Figure 4. A common molt-limit pattern found in certain SY passerines, including Baird's Sparrows. Darker
feathers are formative and paler feathers are juvenal. Illustration: Steve N.G. Howell . .4
Figure 5. Juvenal (white), formative (stippled), and alternate (dark) feathers following the first-prealternate
molt in certain passerines. Illustration: Steve N.G. Howell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Figure 6. Eccentric molt pattern among the primaries and secondaries. Note the replaced formative outer
primaries and inner secondaries contrasting with a block of juvenal feathers in the center of the wing.
Sprague’s Pipits may show this pattern. Illustration: Steve N.G. Howell . .4
Figure 7. Specimens of HY and ASY Sprague’s Pipits (collected in Texas) demonstrating molt limits between
formative and juvenal feathers (HY) and alternate and basic feathers (ASY). Note that the juvenal primary
coverts on HYs are more tapered and worn than the basic primary coverts on ASYs. The difference becomes
more obvious when comparing SYs with ASYs in spring. Photo: Peter Pyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Figure 8. Apparent SY Sprague’s Pipit (from Montana) showing eccentric preformative molt patterns. Note
that the outer four primaries appear fresher, darker, and with darker shafts than the inner five primaries.
The three tertials also appear to have been replaced during the preformative and/or prealternate molts, as
would be typical in individuals replacing the outer four primaries. The left wing showed the same patterns
of replacement. Based on images of the tail, it is difficult to determine whether or not the rectrices were
replaced during the preformative molt. Photo: Janet M. Ruth . .7
Figure 9. Feather generations among wing feathers in an apparent SY Sprague’s Pipit, captured in Montana
in June. J = juvenal, F = formative, and A = alternate feather; red indicates good feathers to sample for
connectivity studies. Photo: Janet M. Ruth . .7
Figure 10. Rectrix shape and condition by age in Sprague’s Pipit specimens (collected in Texas). Note that by
spring the juvenal outer rectrix shown above will become even more worn and frayed in comparison to the
basic outer rectrix. Juvenal rectrices might be replaced during the preformative molt but at what frequency
is unknown (see Figures 11 and 12). Photo: Peter Pyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Figures 11 and 12. Rectrices in two apparent SY Sprague’s Pipits captured for banding in Montana in June.
Both of these individuals showed apparent eccentric patterns in the wings. Many passerines with eccentric
patterns replace all rectrices during the preformative molt but more study is needed (e.g., through
biochemical analysis) to determine rectrix-replacement patterns in this species. Photos: Janet M. Ruth . .8
Figure 13. Grasshopper Sparrow captured for banding in Montana. Note that all wing feathers appear to be
uniform in quality, part of a single generation of either formative or basic feathers. The darker tertials and
List of Figures v
proximal greater coverts represent pseudolimits rather than replaced feathers of a newer generation. Some
of the scapulars as indicated may represent alternate feathers. Photo: Janet M. Ruth . . . . . . . . . . . . . . . . . . 11
Figure 14. Grasshopper Sparrow captured for banding in Montana in June. Note the more worn secondaries
(especially s2-s6) perhaps representing retained juvenal feathers. Photo: Janet M. Ruth . .11
Figure 15. Grasshopper Sparrow captured for banding in Montana in June. Look for darker and fresher
feathers among the back and scapulars that could represent alternate feathers replaced on the wintering
grounds. It would be of interest to compare the biochemical signals from these feathers with those of older,
adjacent feathers. Photo: Janet M. Ruth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Figure 16. Rectrices of a Grasshopper Sparrow captured for banding in Montana in June. The central
rectrices were likely replaced due to accident rather than molt. It would be of interest to sample one of these,
along with one of the other rectrices, to compare locations of development. Photo: Janet M. Ruth . . . . . 11
Figure 17. Specimens of SY and ASY Baird’s Sparrows (collected in Arizona) demonstrating the molt limit
between the formative greater coverts and juvenal primary coverts in the SY, lacking in the ASY. Note the
difference in quality of the primary coverts, and that both individuals had replaced 2-3 inner greater coverts
during the prealternate molt. Photo: Peter Pyle . .13
Figure 18. SY Baird’s Sparrow in Montana in June showing molt limits in the wing. Note that the replaced
formative greater coverts appear fresher and of better quality than the retained juvenal primary coverts
(compare with Figure 19). Note also that the lesser alula appears to be replaced formative whereas the
greater alula appears to be retained juvenal. Due to possible pseudolimits among tertials, it is difficult to tell
whether or not they were replaced at the preformative molt. The juvenal outer primary is also narrow at the
tip. This individual appears not to have replaced any tertials or greater coverts during the prealternate molt
(the darker inner coverts resulting from pseudolimits) but appears to have newer alternate scapulars that
can be used for feather sampling. Photo: Janet M. Ruth . .13
Figure 19. ASY Baird’s Sparrow in Montana in June showing uniformly basic feathers in the wing. In contrast
to the SY in Figure 18, note the darker and higher-quality primary coverts, not contrasting as markedly with
the greater coverts, which are slightly darker in color (naturally) than the primary coverts but uniform in
wear and quality. The outer primaries are also broader and fresher at the tips. No greater coverts or tertials
appear to have been replaced during the prealternate molt but note at least two replaced scapulars near the
joint of the wing. Photo: Janet M. Ruth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Figure 20. Shape and condition of the rectrices in Baird’s Sparrows captured for banding in Montana in June.
Note that the SY appears to have replaced the central rectrices during the preformative and/or prealternate
molt and that the ASY has replaced the right central rectrix during the prealternate molt. Photos: Janet M.
Ruth . .13
Figure 21. Shape and condition of the outer rectrices in SY and ASY Baird’s Sparrows collected in Arizona.
Some individuals show intermediate conditions that would be unreliable to use for aging. Note that the
central rectrices appear to have been replaced during preformative and/or prealternate molts (see Figure
20). Photo: Peter Pyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Figure 22. SY Baird’s Sparrow in Montana in June. Biochemical signals can be obtained for this individual’s
natal site (juvenal secondaries and primaries), wintering site (alternate scapulars, tertials, and inner greater
coverts), and possibly other molting sites (formative greater outer coverts). Photo: Janet M. Ruth . . . . . . . 14
Figure 23. AHY female Chestnut-collared Longspur wing (collected in California) showing uniformly basic
wing feathers. HY/SYs show uniformly juvenal wing feathers throughout the year; thus, molt limits in the
wing cannot be used for aging Chestnut-collared Longspurs, unlike the other species treated here. Note the
broad primary coverts and outer primary tips and the richly colored tertials and greater coverts which, for a
female, combine to indicate an AHY/ASY (Figure 24). Photo: Peter Pyle. . .17
Figure 24. SY female and ASY male Chestnut-collared Longspurs collected on the wintering grounds in
Arizona. Note the more worn wing coverts on the SY, and the lack of white on the inner primary coverts;
within each sex, ASYs have more white than SYs (e.g., compare the SY female here with the AHY female in
Figure 23). Photo: Peter Pyle . .17
Figure 25. Shape and condition of the outer rectrices in Chestnut-collared Longspurs (collected in Texas).
This criterion is reliable throughout the year. Photo: Peter Pyle .17
vii
Acknowledgments
This document resulted from a U.S. Fish and
Wildlife Service, Migratory Bird Office Region 6,
grant to the Institute for Bird Populations (IBP) for
the study of molt. We thank Stephen K. Davis for
sharing images of Sprague’s Pipits at a workshop
in Saskatoon, Saskatchewan, Canada, and David
DeSante (IBP) for support and help administering
the contract. We also thank Steve N.G. Howell for
use of illustrations (Figures 2-6) from Pyle (1997a)
and Maureen Flannery at the California Academy of
Science and Carla Cicero at Museum of Vertebrate
Zoology for allowing examination of specimens. We
thank Michael T. Green, Kevin Kritz, and Maiken
Winter for providing helpful comments on earlier
drafts of this manuscript. This is contribution
number 303 of the Institute for Bird Populations.
Acknowledgments
1
Prairie and grassland habitats in central and
western North America have declined substantially
since settlement by Europeans (Knopf 1994) and
many of the birds and other organisms that inhabit
North American grasslands have experienced steep
declines (Peterjohn and Sauer 1999; Johnson and
Igl 1997; Sauer, Hines, and Fallon 2007). The species
addressed here, Sprague’s Pipit (Anthus spragueii),
Grasshopper (Ammodramus savannarum) and
Baird’s (A. bairdii) sparrows, and Chestnut-collared
Longspurs (Calcarius ornatus), are grassland birds
that are of special conservation concern throughout
their ranges due to declining populations and the
loss of the specific grassland habitats required on
both their breeding and wintering ranges (Knopf
1994, Davis and Sealy 1998, Davis 2003, Davis 2004,
Jones and Dieni 2007).
Population-trend data on grassland birds, while
clearly showing declines, provides no information
on the causes of population declines. Without
demographic information (i.e., productivity and
survivorship), there are no means to determine when
in their life cycle the problems that are creating
these declines are occurring, or to determine to what
extent population trends are driven by factors that
affect birth rates, death rates, or both (DeSante
1995). For migratory birds, population declines
may be driven by factors on breeding grounds,
during migration, and/or on wintering grounds.
Lack of data on productivity and survivorship thus
impedes the formulation of effective management
and conservation strategies to reverse population
declines (DeSante 1992). Furthermore, if deficiencies
in survivorship are revealed, management strategies
may need to address habitats on both breeding and
non-breeding grounds, as well as along migratory
pathways. One technique that helps inform
management strategies is the biochemical analysis of
isotopes and genetic markers, from the sampling of
individual feathers from live birds (Smith et al. 2003,
Pérez and Hobson 2006; Appendix).
Determining demographic parameters and
effectively sampling feathers to reveal connectivity
between breeding and wintering grounds requires
detailed knowledge of molt patterns and age
determination criteria for the target species, in the
hand. For example, productivity, survivorship, and
territory acquisition may all be age-dependent,
with first-year birds showing different patterns and
responses than older birds. In many cases it may
be possible to sample both a feather grown on the
breeding grounds and one grown on the wintering
grounds from a single individual, but knowledge of
age-specific molt patterns, as well as an ability to
recognize different feather generations, is needed
to accomplish such a task. While some information
on molt and aging criteria exists for grassland
passerine species (Pyle 1997a), these species have
been rarely captured during mark-recapture studies
(Jones et al. 2007) and this information thus needs
refining. There is a need for additional resources to
assist field workers in determining molt patterns
and age in captured individuals.
Our objective is to describe molt and aging criteria
for four grassland passerine species with the aid
of digital photographs taken in the field. We hope
that this document will be useful for researchers
studying grassland species through capture and
banding of live individuals on either the breeding or
the wintering grounds. We present a general section
on molt and aging techniques, followed by specific
accounts for the four species treated: Sprague’s
Pipits, Grasshopper and Baird sparrows, and
Chestnut-collared Longspur. We also provide a brief
protocol on collecting feather samples (Appendix).
Molt and Aging by Molt Limits
Molt and plumage terminology used here follow
Humphrey and Parkes (1959) as modified by Howell
et al. (2003), and age terminology follows the
calendar-based system presented by Pyle (1997a).
Preformative and prebasic molts.--Passerines
typically undergo a preformative molt (referred to
as “first prebasic molt” in Pyle 1997a) in their first
summer and fall, and a complete prebasic molt in
July to October following breeding (Figure 1). For
migratory species these molts can occur on the
breeding grounds, on molting grounds away from
breeding or wintering grounds, or on the wintering
grounds; or, they can begin on breeding or molting
grounds, suspend for migration, and complete on
wintering grounds (Pyle 1997a). Which of these
strategies is undertaken can vary both among
species and among individuals of the same species
(Figure 1). In Sprague’s Pipits, the location(s) where
the preformative and prebasic molts occur have yet
to be determined.
The preformative molt undergone by first-cycle,
hatching year/second year (hereafter “HY/SY”),
passerines in July through September (Howell et al.
2003; Figure 1) shows substantial inter- and intra-specific
variation in both extent (varying from partial
Introduction
Introduction
2 Molt and Aging Criteria for Four North American Grassland Passerines
to complete) and in location of occurrence (Pyle
1997a). In some species (including Grasshopper
Sparrows) the preformative molt is typically
complete, and thereafter it becomes difficult or
impossible to separate HY/SY from older after
hatching year/after second year (hereafter “AHY/
ASY”) individuals by plumage-related criteria. In
most passerine species; however, this molt is partial
or incomplete. HY/SYs can be separated from
AHY/ASYs throughout the year by the presence of
“molt limits” (see below for details) within feather
tracts (Figure 2; Pyle 1997a, 1997b; Froehlich
2003), particularly those of the wing (Figure 2),
and sometimes among the rectrices (tail feathers).
Some passerines, particularly those that inhabit
sunny or harsh environments, can also replace outer
primaries during what is known as an eccentric
preformative molt (Pyle 1997a, 1997b; see Figure
6). Although undocumented in the four species
treated here, it appears that at least some Sprague’s
Pipits may undergo such a molt (see species account
below).
Prealternate molt.-- Prealternate molts typically
occur before spring migration on wintering grounds
(February to April). All four species treated here
are reported to have limited or partial prealternate
molts (Figure 1; but see the Grasshopper Sparrow
Figure 1. Molt patterns among four species of grassland passerines during the first and definitive molt cycles.
Solid bars indicate complete molts and dashed bars indicate incomplete, limited, or partial molts. Locations
of where molts occur are indicated above the bars: BG = breeding grounds; WG = wintering grounds.
Figure 2. Tracts of the
Sex, Maids, and Export Processing: Risks and Reasons for Gendered Global Production Networks
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