Phylotranscriptomic analysis uncovers a wealth of tissue inhibitor of metalloproteinases variants in echinoderms

Tissue inhibitors of metalloproteinases (TIMPs) help regulate the extracellular matrix (ECM) in animals, mostly by inhibiting matrix metalloproteinases (MMPs). They are important activators of mutable collagenous tissue (MCT), which have been extensively studied in echinoderms, and the four TIMP copies in humans have been studied for their role in cancer. To understand the evolution of TIMPs, we combined 405 TIMPs from an echinoderm transcriptome dataset built from 41 specimens representing all five classes of echinoderms with variants from protostomes and chordates. We used multiple sequence alignment with various stringencies of alignment quality to cull highly divergent sequences and then conducted phylogenetic analyses using both nucleotide and amino acid sequences. Phylogenetic hypotheses consistently recovered TIMPs as diversifying in the ancestral deuterostome and these early lineages continuing to diversify in echinoderms. The four vertebrate TIMPs diversified from a single copy in the ancestral chordate, all other copies being lost. Consistent with greater MCT needs owing to body wall liquefaction, evisceration, autotomy and reproduction by fission, holothuroids had significantly more TIMPs and higher read depths per contig. Ten cysteine residues, an HPQ binding site and several other residues were conserved in at least 70% of all TIMPs. The conservation of binding sites and the placement of echinoderm TIMPs involved in MCT modification suggest that ECM regulation remains the primary function of TIMP genes, although within this role there are a large number of specialized copies

An Old Lineage of Cyphophthalmi (Opiliones) Discovered on Mindanao Highlights the Need for Biogeographical Research in the Philippines

The arachnid order Opiliones, and the suborder Cyphophthalmi in particular, have recently been used to test biogeographical patterns in Southeast Asia due to their ancient age and extremely low vagility. Here we report the first Cyphophthalmi—two juveniles—known from Mindanao in the southern Philippine Archipelago, and we place them in a molecular phylogeny to test biogeographical hypotheses for their colonization of that island. Five molecular markers were sequenced from one specimen, three from the other, and these sequences were added to a previously completed phylogenetic analysis. The specimens were recovered as members of a clade found almost exclusively on Borneo. Their deep placement within this clade suggests a very old origin and colonization that perhaps involved the mysterious landmass now underlying Mindanao's Zamboanga Peninsula. This species prompts new questions about the abilities of Southeast Asian Cyphophthalmi (Stylocellidae) to disperse and colonize, and it emphasizes how much remains to be understood about the geological history of the Philippines.Organismic and Evolutionary Biolog

Return of chloroquine-sensitive Plasmodium falciparum parasites and emergence of chloroquine-resistant Plasmodium vivax in Ethiopia

BACKGROUND: Increased resistance by Plasmodium falciparum parasites led to the withdrawal of the antimalarial drugs chloroquine and sulphadoxine-pyrimethamine in Ethiopia. Since 2004 artemether-lumefantrine has served to treat uncomplicated P. falciparum malaria. However, increasing reports on delayed parasite clearance to artemisinin opens up a new challenge in anti-malarial therapy. With the complete withdrawal of CQ for the treatment of Plasmodium falciparum malaria, this study assessed the evolution of CQ resistance by investigating the prevalence of mutant alleles in the pfmdr1 and pfcrt genes in P. falciparum and pvmdr1 gene in Plasmodium vivax in Southern and Eastern Ethiopia. METHODS: Of the 1,416 febrile patients attending primary health facilities in Southern Ethiopia, 329 febrile patients positive for P. falciparum or P. vivax were recruited. Similarly of the 1,304 febrile patients from Eastern Ethiopia, 81 febrile patients positive for P. falciparum or P. vivax were included in the study. Of the 410 finger prick blood samples collected from malaria patients, we used direct sequencing to investigate the prevalence of mutations in pfcrt and pfmdr1. This included determining the gene copy number in pfmdr1 in 195 P. falciparum clinical isolates, and mutations in the pvmdr1 locus in 215 P. vivax clinical isolates. RESULTS: The pfcrt K76 CQ-sensitive allele was observed in 84.1% of the investigated P.falciparum clinical isolates. The pfcrt double mutations (K76T and C72S) were observed less than 3%. The pfcrt SVMNT haplotype was also found to be present in clinical isolates from Ethiopia. The pfcrt CVMNK-sensitive haplotypes were frequently observed (95.9%). The pfmdr1 mutation N86Y was observed only in 14.9% compared to 85.1% of the clinical isolates that carried sensitive alleles. Also, the sensitive pfmdr1 Y184 allele was more common, in 94.9% of clinical isolates. None of the investigated P. falciparum clinical isolates carried S1034C, N1042D and D1246Y pfmdr1 polymorphisms. All investigated P. falciparum clinical isolates from Southern and Eastern Ethiopia carried only a single copy of the mutant pfmdr1 gene. CONCLUSION: The study reports for the first time the return of chloroquine sensitive P. falciparum in Ethiopia. These findings support the rationale for the use of CQ-based combination drugs as a possible future alternative

Molecular phylogeny of Indo‐Pacific carpenter ants (Hymenoptera: Formicidae, Camponotus) reveals waves of dispersal and colonization from diverse source areas

Ants that resemble Camponotus maculatus (Fabricius, 1782) present an opportunity to test the hypothesis that the origin of the Pacific island fauna was primarily New Guinea, the Philippines, and the Indo‐Malay archipelago (collectively known as Malesia). We sequenced two mitochondrial and four nuclear markers from 146 specimens from Pacific islands, Australia, and Malesia. We also added 211 specimens representing a larger worldwide sample and performed a series of phylogenetic analyses and ancestral area reconstructions. Results indicate that the Pacific members of this group comprise several robust clades that have distinctly different biogeographical histories, and they suggest an important role for Australia as a source of Pacific colonizations. Malesian areas were recovered mostly in derived positions, and one lineage appears to be Neotropical. Phylogenetic hypotheses indicate that the orange, pan‐Pacific form commonly identified as C. chloroticus Emery 1897 actually consists of two distantly related lineages. Also, the lineage on Hawaiʻi, which has been called C. variegatus (Smith, 1858), appears to be closely related to C. tortuganus Emery, 1895 in Florida and other lineages in the New World. In Micronesia and Polynesia the C. chloroticus‐like species support predictions of the taxon‐cycle hypothesis and could be candidates for human‐mediated dispersal.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/1/cla12099-sup-0002-FigureS2.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/2/cla12099-sup-0003-FigureS3.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/3/cla12099-sup-0001-FigureS1.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/4/cla12099-sup-0004-FigureS4.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/5/cla12099-sup-0005-FigureS5.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/6/cla12099-sup-0006-FigureS6.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/112260/7/cla12099.pd

An ant genus-group ( Prenolepis ) illuminates the biogeography and drivers of insect diversification in the Indo-Pacific

The Malay Archipelago and the tropical South Pacific (hereafter the Indo-Pacific region) are considered biodiversity hotspots, yet a general understanding of the origins and diversification of species-rich groups in the region remains elusive. We aimed to test hypotheses for the evolutionary processes driving insect species diversity in the Indo-Pacific using a higher-level and comprehensive phylogenetic hypothesis for an ant clade consisting of seven genera. We estimated divergence times and reconstructed the biogeographical history of ant species in the Prenolepis genus-group (Formicidae: Formicinae: Lasiini). We used a fossil-calibrated phylogeny to infer ancestral geographical ranges utilizing a biogeographic model that includes founder-event speciation. Ancestral state reconstructions of the ants\u27 ecological preferences, and diversification rates were estimated for selected Indo-Pacific clades. Overall, we report that faunal interchange between Asia and Australia has occurred since at least 20–25 Ma, and early dispersal to the Fijian Basin happened during the early and mid-Miocene (ca. 10–20 Ma). Differences in diversification rates across Indo-Pacific clades may be related to ecological preference breadth, which in turn may have facilitated geographical range expansions. Ancient dispersal routes suggested by our results agree with the palaeogeography of the region. For this particular group of ants, the rapid orogenesis in New Guinea and possibly subsequent ecological shifts may have promoted their rapid diversification and widespread distribution across the Indo-Pacific

Camponotus erythrocephalus Clouse, 2007, n. sp.

<i>Camponotus erythrocephalus</i> n. sp. <p>Plate 1 D–F</p> <p>HOLOTYPE WORKER: TL 3.95, HL 1.02, HW 0.95, CI 93, SL 1.30, SI 137, PW 0.73, ML 0.38. Mandible outer margin strongly curving to a sharp apical tooth, the apex parallel to the anterior clypeal margin. (Holotype with material in mandibles, so mandibles and anterior clypeus described below from paratypes.) Median clypeus tapering and dipping away from the vertex centrally, antennal insertions separated from clypeus by a distance slightly more than the distance from nearest clypeal margin to the clypeal midpoint. In front view, head tapering slightly toward the mandibles, vertex rounded, eyes at upper corners. Bottom margin of eyes located above halfway point of the head and slightly above level of the antennal insertions; their inner margin slightly less than halfway from frontal lobes to sides of the head, their outline breaking the outline of the sides of the head. Antennae 12-segmented, the scape extending beyond the vertex by more than half its length. Mesosoma broad and shallow in profile, sloping rather evenly from the anterior pronotum to the posterior propodeum, a slight depression at the metanotal groove and an even slighter one at the promesonotal suture, propodeal declivity distinct but slight. Petiole about 1.5 times longer than tall, its anterior edge rising only about the same distance as the length of the peduncle, then sloping up for a distance slightly less than the height of the posterior edge. Color: Reddish orange head and mesosoma contrasting against a mostly black gaster. Head even in color, mandibles same shade as the remainder of the head. Mesosoma, legs, and petiole same color as head, although coxae slightly darker. Gaster not entirely black but with four grey stripes made by whitishclear strip along posterior of each gastral tergite; strip is slightly wider than maximum width of the scape. Anterior gastral tergites approach color of mesosoma centrally. Pilosity: Head dominated by recumbent silver hairs interspersed with long, standing hairs on frons and central clypeus. Standing hairs vary in height, but pair closest to vertex as long as maximum width of eyes. Mesosoma with even silver sheen created by recumbent hairs, propodeum with two pairs of long, standing hairs. Petiolar node with four long, backward-pointing hairs arranged in two pairs, one on each side of its highest point; inner hairs slightly shorter than outside ones. Each gastral tergite with about ten long, silver hairs immediately anterior to posterior grey strip. In addition, two adjacent pairs of long hairs lie in line along middle of each tergite. Sculpturing: Head with slight puncturing and body with shallow whorls, but mostly shining.</p> <p>PARATYPE WORKERS: TL 4.50–5.15, HL 1.23–1.28, HW 1.03–1.13, CI 81–93, SL 1.37–1.42, SI 121– 137, PW 0.82–0.93, ML 0.40–0.47. Five teeth on the masticatory margin visible when the mandibles are closed, declining in size from the apex. Clypeal margin almost straight but curving out slightly. Scapes extending beyond the vertex by a distance less than half their length.</p> <p> <b>Similar species:</b> The flattened petiole, monochromatic reddish anterior and contrasting black gaster appear unique in this species. Among unidentified collections from Borneo, only two species have a resemblance, but the one closest in coloration has a tall petiole, and the one similar in shape is monochromatic black and overall too narrow. Given that it has only been collected on Yap Island in Micronesia, it appears to be a poor disperser and could easily be an island endemic.</p> <p> <b>Etymology:</b> <i>Erythrocephalus</i> is Latin for “red-headed,” which is the distinguishing feature of this species.</p> <p> <b>Type locality:</b> Yap Island, Yap State, Federated States of Micronesia</p> <p> <b>Type series:</b> Holotype worker: FSM Yap: Yaptown, Yap I., Mt. Matade (“No. 1087,” Townes, 12-VII- 1946, NMNH). Paratype workers (2): FSM Yap: Yap I. (Goss, “Jul–Aug” 1950)</p>Published as part of <i>Clouse, Ronald M., 2007, New ants (Hymenoptera: Formicidae) from Micronesia, pp. 1-19 in Zootaxa 1475</i> on page 6, DOI: <a href="http://zenodo.org/record/273767">10.5281/zenodo.273767</a&gt

Camponotus eperiamorum Clouse, 2007, n. sp.

Camponotus eperiamorum n. sp. Plate 1 A–C HOLOTYPE WORKER: TL 5.60, HL 2.15, HW 1.70, CI 79, SL 2.00, SI 118, PW 1.15, ML 0.95. Mandible outer margin gently curved to an apex of about 75 degrees, the masticatory margin straight in front view, leading to an angle of 100 degrees and a basal margin about half the length of the masticatory margin. Masticatory margin with six teeth, which gradually diminish in size from the apex (the basal tooth in some specimens broad or with a slight double point). Diastema between basal tooth and angle equal to half the width of basal tooth. Clypeus continuing anteriorly past mandibular insertions a distance slightly less than length of apical tooth, then straight across. Median clypeus curved away from vertex, antennal insertions separated from clypeus by a distance slightly more than distance from nearest clypeal margin to clypeal midpoint. Head longer than wide. In frontal view, eyes located vertically halfway between posterior clypeus and vertex; horizontally, inner margins halfway between frontal lobes and sides of the head, their width reaching halfway to the lateral edge of the head in larger workers and all the way in smaller ones. Antennae 12 -segmented. Scape extending beyond the vertex by 2 / 5 its length. Mesosoma in profile gently sloping from anterior pronotum to dorsal propodeum, with moderate propodeal declivity. Petiolar node moderate height, anterior face half the height of posterior face and parallel. Color: Mesosoma yellow, gaster glossy black, head mostly deep orangebrown. Each gastral tergite with clear strip along posterior fifth. Head abruptly becomes same color as dorsal pronotum at vertex; border separating the coloration of vertex from remainder of the head located from the mid-vertex by a distance about as wide as width of the scape, dropping down to level of the eyes laterally. Mandibles darker than head and contrasting with clypeus. Clypeus similar in color to vertex, especially toward mandibles. Teeth of the mandibles, scrobes, sutures, and joints on the head darker than surrounding cuticle. Pilosity: On head, frontal area lateral to the eyes lacking long, standing hairs; with layer of small, recumbent, light hairs all over head; long, standing hairs numerous on front and back, frontal ones starting at clypeus and continuing to vertex, becoming less numerous but longer. Mesosoma with three pairs of long, standing hairs clustered on pronotum, two pairs clustered on mesonotum, and ten stout hairs of various lengths scattered on the propodeum. Each gastral tergite with 6–9 rather evenly spaced, short, standing hairs immediately before clear strip along posterior edge. Just after end of each tergite a row of four longer hairs on next tergite. Sculpturing: Surface smooth and shining, although not glossy. PARATYPE WORKERS: TL 5.10–6.50, HL 1.55–2.55, HW 1.05–2.30, CI 68–90, SL 1.95–2.20, SI 88– 200, PW 0.85–1.30, ML 0.60 –1.00. As in the holotype except head approaching square in larger workers, scape extending beyond the vertex by a range of 1 / 2 to 1 / 6 scape length in smaller to larger workers, some workers darker but retaining same contrast between body parts, area lacking long hairs lateral to the eyes extending around to the lateral back of head in smaller workers. PARATYPE QUEENS: TL 8.50–9.30, HL 2.20–2.40, HW 2.05–2.15, CI 88–93, SL 2.00– 2.20, SI 95– 107, PW 1.85–2.10, ML 0.90–1.10. Similar to workers except for larger mesosoma and sclerites associated with wings. Also, eyes larger and located more laterally than in major workers, their profile meeting the sides of the head in frontal view. Heads more square than those of largest workers. Mesosoma large, three times wider than workers and swollen between first and middle coxae. Metanotal tergite dark brown to black, contrasting sharply with yellow-orange mesosoma. PARATYPE MALE: TL 6.00, HL 1.10, HW 1.20, CI 109, SL 1.40, SI 117, PW 1.45, ML 0.45. Mandible edentate, curving to a point at the apex, angle rounded, basal margin twice length of masticatory margin. Posterior clypeus bilobed, curving down from each side to a point medially. Frontal lobes reduced, interrupted by torulus. Compound eyes large, ocelli large and elevated. Antennae 13 -segmented. Mesosoma large, pronotum and mesosternum swollen. Petiolar node short, anterior and posterior faces not distinctly parallel. Coloration of mandibles, clypeus, and between the antennal insertions light orange to yellow; rest of head dark orange ranging to near black between the ocelli. Metanotal tergite dark brown, as with female. Similar species: Among species in Micronesia, Camponotus eperiamorum n. sp. is most similar to C. chloroticus Emery 1897, and it is thus a member of the difficult maculatus -group in the Pacific. (Camponotus maculatus was originally described by Fabricius in 1782, but see Wilson and Taylor [1967] and Clouse [in press] for more history.) Morphologically, the only difference I can find between eperiamorum and chloroticus is the dearth of long hairs on the sides of the head in eperiamorum. The coloration, however, is quite distinct and makes eperiamorum easy to identify in the field. Another close relative of chloroticus in the Pacific is C. navigator (Wilson & Taylor 1967) in Polynesia. However, navigator differs from chloroticus (and thus, generally, eperiamorum) in several small but distinct morphological features, and its coloration (overall reddish to blackish brown) is very different from both chloroticus and eperiamorum. The bicoloration of eperiamorum resembles some forms in the C. maculatus group, especially C. arrogans (Smith 1858). However, eperiamorum is unique in having an extremely light yellow mesosoma and displaying remarkable consistency in its coloration among various collections. Camponotus arrogans is more reddish, and its head is uniform in coloration, different from eperiamorum, which lacks red or orange and has a light clypeus and vertex. Moreover, other forms, including arrogans, have significant pilosity on the sides of the head, unlike eperiamorum, and they often have light markings on the gaster, absent in eperiamorum. Finally, eperiamorum appears to be a native Pohnpeian species from its habitat of high- to mid-elevation native forest. Etymology: This species is named in honor of the Eperiam family of Einpein Village in Kitti Municipality on Pohnpei Island. Emensio Eperiam has had a long career in environmental protection, historical preservation, and tourism on Pohnpei. His much-loved, late wife, Mercedes, and their family demonstrated extreme hospitality and generosity to those of us who wanted to hike to the interior. His sons Abram, Casandro, and Paulo guided me to remote areas and kept me from becoming lost while I collected. Type locality: Pohnpei Island, Pohnpei State, Federated States of Micronesia, in mid-elevation forest. Type series: Holotype worker and 2 paratype workers: FSM Pohnpei: Pohnpei I., Mt. Nankep [“Mt. Delennankap” on label] (1700–2000 ft., forest plants, Townes, 10 -VIII- 1946, NMNH). Other paratypes (85 workers, 4 queens, 1 male): FSM Pohnpei: Pohnpei I., Kitti (on dead fern branch in high-elevation sakau clearing, Clouse, 24 -III- 2000), Lehnpeinpohn Waterfall (on road to falls, under dead leaves on ivory nut tree, Okihiro and Clouse, 26 -XI- 1995), Mahnd (along river above village, elevation 200 m, large nest inside dead ivory nut branch, Clouse, 29 -X- 1994), Mahnd (0.5 mile up river from village, large nest in dead tree fern branch leaning on ivory nut tree, Clouse, 29 -X- 1995), Malen Pahnpe (above Keprohi Falls at elevation 350 m, on ivory nut along river, Clouse, 9 -VII- 1994), to Mt. Nahnalaud (upland forest camp near river at elevation 300 m, carrying food up tree, Clouse, 5 -III- 1995), to Mt. Nahnalaud from Kitti (walking on dead log at elevation 300 m, Clouse, 8 -IX- 1995), Mt. Nankep (“ 1660,” forest plants, Townes, 8 -II- 1946, NMNH), Awak (alates at light at residence, Okihiro, 9 -VI- 1996) Other specimens examined: FSM Pohnpei: Pohnpei I., Nankep (1800 ft., [three specimens on monocot midribs—possibly elephant grass, Pennisetum purpureum Schumach—clenching it with mandibles, covered with dried hyphae and with a ~ 2.5 cm fruiting body coming out from behind each head], Townes, 13 -VIII- 1946, NMNH)Published as part of Clouse, Ronald M., 2007, New ants (Hymenoptera: Formicidae) from Micronesia, pp. 1-19 in Zootaxa 1475 on pages 3-5, DOI: 10.5281/zenodo.27376

Vollenhovia kaselela Clouse, 2007, n. sp.

Vollenhovia kaselela n. sp. Plate 5 HOLOTYPE WORKER: TL 2.00, HL 0.54, HW 0.45, CI 84, SL 0.31, SI 69, PW 0.35, ML 0.25, PL 0.21, DPW 0.17, PWI 81, PPL 0.20, PPW 0.19, PPWI 95. Mandible outer margin straight, curved at apical tip. Masticatory margin with six teeth, apical and penultimate teeth sharp, remaining four teeth blunter, decreasing in size basally. Head roughly rectangular, tapering slightly toward mandibles, median vertex concave. Eyes lateral, upper margin of eyes at mid-level of head. Antennae 12 -segmented with indistinct, three-segmented club; first segment of club three times as large as previous, second segment of club 80 % larger, and terminal antennal segment twice as large as penultimate. Frontal lobes not hiding torulus completely, stopping at insertions. Mesosoma nearly flat in profile, trace of metanotal groove, gently sloping at propodeal declivity but with small latero-dorsal corners. Propodeal lobes extending nearly halfway up declivity and posteriorly the same distance. Petiole (not including keel) roughly cylindrical, anterior node rising vertically, with small tooth in profile at location of spiracle then sloping up to dorsal surface; tapered dorsally, ridged around posterior socket. Peduncle indistinct. Ventral petiolar keel constituting one fifth total petiolar height, anterior edge vertical, dorsal edge sloping up at about 45 degrees to point just anterior to postpetiolar insertion; lower extreme of keel translucent. Postpetiole globular, ventral edge concave, with lip ventrally at petiolar juncture. Gaster elliptical, 80 % covered by first tergite. Color: Most of head, mesosoma, petiole, postpetiole, gaster, forecoxae, and femora blackish brown; mandibles, clypeus and anterior head around mandibular insertions, and remain- der of legs orange-brown. Pilosity: Mostly covered with mixture of long, straight, standing hairs and short hairs curved back to the body surface. Clypeus with two long, straight hairs immediately below antennal insertions, pointing toward midline. Hairs on face shorter and more depressed than remainder of body, pointed toward vertical midline. Pilosity on dorsal mesosoma as with head, low and curved toward center, but mixed with a few very long, standing hairs along the dorso-lateral edge. Hairs less numerous but longer on dorsal petiole and postpetiole. Lateral propodeum and mesepisternum as well as lateral and anterior faces of petiole hairless. Dorsal and ventral gaster with curved, short hairs and long, straight ones mixed. Mandibles, antennae, lateral pronotum, and legs with short, curved hairs. Sculpturing: Head striae starting immediately above median clypeus, gena with distinct striae but no punctures or hairs. Small smooth patches lacking on dorsal pronotum, mesonotum, or propodeum. Declivity and entire dorsal petiole and postpetiole smooth. PARATYPE WORKER: TL 1.90, HL 0.55, HW 0.48, CI 86, SL 0.31, SI 66, PW 0.38, ML 0.26, PL 0.23, DPW 0.18, PWI 78, PPL 0.20, PPW 0.20, PPWI 100. Similar species: The only other small Vollenhovia species with such extensive punctures and striations on the head is an unidentified specimen from Chuuk Island (Clouse, in press); however, kaselela is easily distinguished from it by its larger size and proportionately wider petiolar node. Vollenhovia kaselela is also darker, has narrower striations on the gena, more puncturing on the pronotum, and less sculpturing on the dorsal petiolar node than this Chuukese specimen. The Melanesian species V. subtilis Emery 1887 also has striate genae but in addition to being redder in color and slightly larger than kaselela, is less punctured, being smooth just anterior of the propodeal spiracle and on the dorsal propodeum. The blanket of regular punctures on Vol le n - hovia kaselela is approached in the similarly sized V. e m e r y i Wheeler 1906 (especially its subspecies chosenica Wheeler 1928) and V. banksi Forel 1910 b subspecies kuchingensis Wheeler 1919 from Borneo, but they do not have striate gena. Etymology: Kaselel means “beautiful” or “perfect,” in Pohnpeian. Type locality: Pohnpei Island, Federated States of Micronesia Type series: Holotype worker: FSM Pohnpei: Pohnpei I., Nett Municipality, Nanpil (Dybas, 25 -II- 1948). Paratype worker (1): FSM Pohnpei: Pohnpei I., PATS (walking on rotten log, collected by hand, 21 -III- 2000, Clouse)Published as part of Clouse, Ronald M., 2007, New ants (Hymenoptera: Formicidae) from Micronesia, pp. 1-19 in Zootaxa 1475 on pages 13-15, DOI: 10.5281/zenodo.27376

Pheidole recondita Clouse, 2007, n. sp.

Pheidole recondita n. sp. Plate 4 HOLOTYPE MAJOR WORKER: TL 2.33, HL 1.02, HW 0.98, CI 97, SL 0.40, SI 41, PW 0.55, ML 0.45. Outer mandibles curved at right angles, apical margins parallel to each other and perpendicular to anterior clypeus. Apical margins worn in holotype but paratype with teeth, described below. Anterior clypeus concave centrally, median clypeus extending to level of antennal insertions, depressed from surface at highest point. Head mostly square, bilobed at vertex. Eyes lateral, at level slightly higher than halfway up scapes. Frontal lobes hiding antennal insertions, scrobes distinct, extending slightly beyond reach of scapes. Antennae 13 - segmented, three-segmented club equal in length to remaining funiculus, terminal segment same length as preceding two combined. Promesonotum bulbous in profile, meeting dorsal propodeum at right angle, extended laterally. Propodeal spines as long as height of posterior propodeum, spiracle round and prominent. Petiole angled up from long peduncle, anterior height equal to dorsal length of peduncle. Postpetiole slightly shorter than petiole but expanded to greater width laterally. Color: Overall blackish brown fading to orange at legs and distal scapes through funiculus. Especially blackish at mandibular margins, head along mandibular insertions, proximal scapes, mesepisternum, metepisternum, and posterior gastral tergites. Pilosity: Front of head, vertex, dorsal promesonotum, dorsal petiole and postpetiole, and dorsal gaster with scattered, long, curved, light hairs, especially long and curved on the promesonotum. Sculpturing: Distinguished by extensive puncturing on head, mesosoma, petiole, and postpetiole, even among striae and reticulations; puncturing missing from clypeus, between antennal insertions, and patches on the lateral pronotum, mes- and metepisterna. Reticulate sculpturing at vertex sharp and wide, becoming more linear and losing intermingled punctures at top of scrobes, and ending as linear striae from the middle of the scrobes to the median clypeus. Reticulations continuing down sides of head, becoming more linear around eyes but less distinct among punctures between eyes and antennal insertions, ending in fine linear sculpturing in front of mandibular insertions. Scrobes lined with punctures only. Clypeus and mandibles smooth. Promesonotum covered in loose rugae mixed with punctures, becoming predominated by punctures laterally and between propodeal spines. Lateral pronotum directly under lateral pronotal extensions, mesepisternum, and metepisternum each with glossy area free from punctures and rugae. Petiole and postpetiole with weaker sculpturing, mostly seen as punctures on dorso-lateral peduncle. Gaster smooth but not glossy. PARATYPE MAJOR WORKERS: TL 2.32–2.80, HL 0.87–1.12, HW 0.87–1.07, CI 92–100, SL 0.35– 0.45, SI 40–47, PW 0.45–0.58, ML 0.45–0.53. As with holotype major worker except: Mandibles not as worn, with two apical teeth and hint of a basal tooth, with a gently concave, edentate surface between them. Apical tooth larger than penultimate. CI increases and SI decreases in smaller individuals, seen in paratype from same collection as holotype and paratypes from Rota. Pohnpeian paratypes largest. More recently collected specimens more blackish than holotype and paratype from same collection as holotype. Punctures visible between linear striae between antennal insertions. PARATYPE MINOR WORKERS: TL 1.00– 1.16, HL 0.40–0.46, HW 0.40–0.45 CI 98–105, SL 0.34– 0.43, SI 83–96, PW 0.26–0.28, ML 0.22–0.25. Mandible outer margin angled straight in to center, ending with a long, backward-curving apical tooth, followed by a second tooth of about half length, then six irregular and often indistinct denticles along apical margin. Frontal lobes not hiding torulus, scrobes absent. Eyes lateral and bulging, slightly below midlevel of head. Vertex concave. Scapes extending beyond vertex by distance less than first segment of funiculus. Mesosoma humped in profile, promesonotum meeting dorsal propodeum at 120 degree angle. Propodeal spines similar in length to height of posterior propodeum and length of petiolar peduncle. Peduncle straight dorsally, bulging ventrally. Petiolar node taller than postpetiole, which is expanded gently laterally. Color: Body blackish brown to brownish black, becoming orange at funiculus, mandibles, and legs. Pilosity: Many fewer hairs than majors, except for clypeus, mandibles, and antennae, which have numerous, long, standing, silver hairs. Sculpturing: Head (including clypeus), mesosoma, and dorsolateral peduncle covered with deep, numerous, and evenly distributed punctures. Hints of linear sculpturing on clypeus, between eyes and antennal insertions, and bordering mesepisternum. Similar species: Pheidole recondita n. sp. is one of three extremely small, punctured Pheidole in Micronesia and Melanesia, the other two being P. nindi Mann 1919 and P. philemon Forel 1910 a. Although the three majors are quite easily distinguished, the minors are extremely similar and difficult (but not impossible) to distinguish without associated majors. Pheidole recondita n. sp. and philemon majors are distinguished from nindi by their long frontal lobes: in nindi, linear sculpturing on the frons gradually blends with punctures that lie under the scapes, whereas in recondita and philemon, the two types of sculpturing are distinct and separated by a continuation of the frontal lobes along the length of the scapes. Pheidole recondita and philemon majors are distinguished from each other by the sculpturing of the mesosoma, which is generally more punctate in recondita. Although punctured, philemon is mostly covered with reticulate sculpturing or rugae which become fine transverse ridges on the dorsal pronotum and fade to smooth surface along the lateral mesosoma; recondita has irregular rugae that continue down the sides of the mesosoma, except for distinct, small patches on the lateral pronotum, mes- and metepisterna. A character that is extremely useful to distinguish these three species—even the minors—is their coloration: nindi ranges from light orange to reddish brown, recondita ranges from blackish orange to blackish brown (always with a blackish or ashy appearance), and philemon is bicolored, with a dark brown gaster and ventral petiole and postpetiole and light orange head, mesosoma, and dorsal petiole and postpetiole. Other than color, the minors can be distinguished as follows: nindi and recondita have straight, triangular propodeal spines, while philemon has spines which curve slightly downward; philemon is smoother on the dorsal promesonotum than the other two species, and it has slight transverse rugae on the dorsal mesonotum (nindi and recondita are both heavily punctate over the entire mesonotum); and the dorsal propodeum in recondita is short and convex relative to nindi, which has a relatively long, flat propodeum posterior to a slight anterior rise. Etymology: Recondita is Latin for “mysterious,” as is this uncommon little ant. The fact that it has been collected from three different areas in Micronesia (Kosrae, Pohnpei, and the Marianas) suggests that it is a good disperser, but its absence from collections outside Micronesia make it one of only two species that currently stand as Micronesian endemics found on more than one island. The other one, Metapone truki Smith, M.R., 1953 from Chuuk and Palau, is currently believed to be a junior synonym of a described species widely found on New Guinea (R. W. Taylor, pers. comm.), thus making P. recondita ’s distribution potentially unique. Type locality: Micronesia, from Guam Island and Rota Island in the Mariana archipelago, and Pohnpei Island of the Caroline Islands. (Non-types have been examined from Kosrae Island, also in the Carolines.) Type series: Holotype major worker: Mariana Is.: Guam I., Yigo (Swezey, 18 -X- 1936). Paratype workers (4 majors associated with 12 minors): FSM, Pohnpei: Pohnpei I.: Mt. Nahnalaud Cave (under moss on tree around camp, collected by hand, Clouse, 24 -III- 2000); Mariana Is.: Guam I., Yigo (Swezey, 18 -X- 1936); Rota I. (litter, shaded valley, Bourquin, 15 -XI- 2002); Collections of minors examined (not made paratypes): FSM Kosrae I.: Mt. Mutunte (580 m, duff from forest floor; KU 58 B, Clarke, 11 -II- 1953); FSM Pohnpei: Pohnpei I., Mt. Nahnalaud (collected by Berlese funnel from moss, Clouse, 10 -IX- 2000); Mariana Is.: Anatahan I. (pitfall, mid-altitude forest, Bourquin, 2 -IV- 2002); Anatahan I. (pitfall, mid altitude forest, Bourquin, 26 -IV- 2002)Published as part of Clouse, Ronald M., 2007, New ants (Hymenoptera: Formicidae) from Micronesia, pp. 1-19 in Zootaxa 1475 on pages 10-13, DOI: 10.5281/zenodo.27376