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Neal Woodman

Publications and source records attributed to Neal Woodman.

At least 19 recordsLinked to original sources

Records of fleas (Siphonaptera) from Delaware

We present an annotated checklist of fleas (Siphonaptera) known to occur in the state of Delaware based on an examination of Siphonaptera collections at the University of Delaware and the Carnegie Museum of Natural History, as well as new specimens of fleas we collected from wildlife, other hosts, and tick flags. We review published records and compile them herein with our new records, which include 3 species previously unreported from Delaware. With these additions, there are now 18 flea species from 19 avian and mammalian hosts documented from Delaware.

Delaware

Skeletal indicators of locomotor adaptations in shrews

The Soricidae (Mammalia: Eulypotyphla) comprises more than 450 species inhabiting a variety of habitats on five continents. As a family, shrews employ a variety of locomotor modes that incorporate ambulatory, fossorial, aquatic, and scansorial behaviors, illustrating an ability to exploit a variety of natural substrates and their associated resources. In this study, the association of skeletal morphology and three of the dominant locomotor modes in the family—ambulatory, semi-fossorial, and semi-aquatic behaviors—was investigated in up to 52 species of 12 genera representing all three subfamilies of Soricidae. From skeletal measures, 34 morphological indices were calculated, most of which have been used previously to characterize substrate use among shrews, rodents, and other mammals, and analyzed for their individual effectiveness for discriminating the three locomotory modes. To assess their effectiveness in combination, subsets of locomotor indices were analyzed using 1) mean percentile ranks, 2) the first principal component from principal components analysis, and 3) plots and classifications from discriminant function analyses. In general, the three methods effectively identified and grouped the three locomotor modes and identified smaller subsets. Additional analyses were then used to classify the locomotor behaviors of five species whose locomotor modes were unknown or ambiguous. The analyses reinforce and broaden the scope of a previously identified observation of the wide range of grades of morphological variation that may permit an equally diverse range of locomotor abilities among the Soricidae.

Therya

Two new species of small-eared shrews of the Genus Cryptotis Pomel, 1848, from the Colombian Andes (Mammalia: Eulipotyphla: Soricidae)

Shrews (Mammalia: Eulipotyphla: Soricidae) reach the southern limit of their New World distribution in the Andes and eastern coastal highlands of northern South America. South of Honduras, the family is represented only by species of the genus Cryptotis Pomel, 1848. In South America, soricids are restricted to moist, high-elevation environments above 1000 m, and their distribution appears to be discontinuous. Study of specimens from a previous gap in the known geographical range of shrews in the Central Cordillera of southwestern Colombia reveals the presence of two unique populations that are distinguishable from each other and their congeners by a combination of morphological and morphometrical characters. They are described herein as, Cryptotis huttereri, n. sp. and Cryptotis andinus, n. sp. Both species are members of the Cryptotis thomasi group, one of five species groups of small-eared shrews defined partly on the basis of postcranial morphology and potential locomotor behavior. Although species in the C. thomasi group share similar postcranial architecture, as exemplified by the morphology of the forelimb, the group appears to be polyphyletic, implying convergence in locomotor behavior, possibly one uniquely adapted for Andean-type montane habitats. Recognition of C. huttereri and C. andinus brings the total number of known South American soricids to 19 species, with 11 species occurring in Colombia. Of those, seven species are endemic to that country.

Annals of the Carnegie Museum

Environmental implications of Ptolemaic Period rodents and shrews from the Sacred Falcon Necropolis at Quesna, Egypt (Mammalia: Muridae and Soricidae)

Background Assemblages of mummified and preserved animals in necropoleis of Ptolemaic Period Egypt (ca. 332–30 BC) document some aspects of the ceremonial and religious practices of the ancient Egyptians, but study of these animal remains can also provide insight into the local environments in which the animals and humans lived. Results Excavations of the Sacred Falcon Necropolis at Quesna in the Nile Delta have yielded many thousands of animal remains, mostly of raptors, but also of a lesser number of small, wild mammals. Among the latter, we identified four species of murid rodents (Rodentia: Muridae) and five species of shrews (Eulipotyphla: Soricidae). The soricids are of particular interest because they represent a more diverse assemblage of species than occurs in the delta today. They include one species, Crocidura gueldenstaedtii (Pallas, 1811), that no longer occurs in the delta and another, C. fulvastra (Sundevall, 1843), that is now extirpated from Egypt. Conclusions The coexistence of this diverse small mammal community suggests that a greater availability and variety of mesic habitats were present during the Ptolemaic Period than occur there now. The local mammal faunas recovered at Quesna and other well-studied ancient Egyptian sites together provide evidence of a richer, more complex regional environment along the Nile Valley. They also provide important insight regarding the biogeography of the individual species comprising the faunas and about the extent of faunal turnover since the Ptolemaic Period.

BMC Ecology and Evolution

Taxonomic boundaries in Lesser Treeshrews (Scandentia, Tupaiidae: Tupaia minor)

The Lesser Treeshrew, Tupaia minor Günther, 1876 , is a small mammal from Southeast Asia with four currently recognized subspecies: T. m. minor from Borneo; T. m. malaccana from the Malay Peninsula; T. m. humeralis from Sumatra; and T. m. sincepis from Singkep Island and Lingga Island. A fifth subspecies, T. m. caedis , was previously synonymized with T. m. minor ; it was thought to occur in northern Borneo and on the nearby islands of Banggi and Balambangan. These subspecies were originally differentiated based on pelage color, a plastic feature that has proven to be an unreliable indicator of taxonomic boundaries in treeshrews and other mammals. To explore infraspecific variation among T. minor populations across the Malay Peninsula, Borneo, Sumatra, and smaller islands, we conducted multivariate analyses of morphometric data collected from the hands and skulls of museum specimens. Principal component and discriminant function analyses reveal limited differentiation in manus and skull proportions among populations of T. minor from different islands. We find no morphometric support for the recognition of the four allopatric subspecies and no support for the recognition of T. m. caedis as a separate subspecies on Borneo.

Borneo, Sumatra

Whence Orycteropus? The correct authorship and date for the generic name of the aardvark (Mammalia, Tubulidentata, Orycteropodidae)

All else being equal, the principle of priority in zoological taxonomic nomenclature gives precedence to the earliest name for a particular taxon. Determining the origin of some late eighteenth- and early nineteenth-century taxonomic names, however, can be vexing, particularly when the history of a name was never completely documented in contemporary synonymies. The authorship and date for Orycteropus Étienne Geoffroy Saint-Hilaire, 1796: 102, the genus-group name for the African aardvark, Orycteropus afer (Pallas, 1766), has been variously ascribed to at least four authors other than É. Geoffroy Saint-Hilaire. Using digitally imaged publications now available in a variety of internet-accessible libraries, I traced the comprehensive history of the name and show how and, to some extent, why its origin became obscured. É. Geoffroy Saint-Hilaire’s original description was re-published twice, most likely to make the description more widely available. Rather than reinforce his authorship for the name, however, the surprising consequence of the multiple publications was to cast doubt on it.

Bionomina

The relevance of a type locality: The case of Mephitis interrupta Rafinesque, 1820 (Carnivora: Mephitidae)

For more than 130 years, the type locality of the Plains Spotted Skunk, Spilogale putorius interrupta (Rafinesque, 1820) has been accepted to be along the upper Missouri River. The species’ description was based on a specimen observed by Constantine S. Rafinesque during his 1818 exploration of the Ohio River Valley, but Rafinesque never ventured into the animal’s geographic range west of the Mississippi River, calling into question the type locality and, therefore, the identity of the taxon. We reconstruct Rafinesque’s itinerary from his notes, publications, and correspondence and determine that Rafinesque probably observed the specimen on 20 September in Middletown, Kentucky, while traveling between Louisville and Lexington. He spent the day with John Bradbury, who participated in the 1811 Astor expedition up the Missouri River. On 1 April 1811, Bradbury collected the skin of a skunk, and evidence suggests that it was this skin that Rafinesque described. The type specimen of the Plains Spotted Skunk was obtained on the Missouri River flood plain in southern Chariton County or northern Saline County, Missouri, and this area should be considered the type locality for M. interrupta .

Illinois, Indiana, Kentucky, Missouri, New York, O

Ecogeographic variation and taxonomic boundaries in Large Treeshrews (Scandentia, Tupaiidae: Tupaia tana Raffles, 1821) from Southeast Asia

The Large Treeshrew, Tupaia tana Raffles, 1821, is a small mammal (~205 g) from Southeast Asia with a complicated taxonomic history. Currently, 15 subspecies are recognized from Borneo, Sumatra, and smaller islands, and many were originally differentiated based on minor pelage differences and small sample sizes. We explored intraspecific variation in T. tana using quantitative osteological data obtained from the hands and skulls of museum specimens. Multivariate analyses reveal extensive overlap among T. tana populations in morphospace, indicating that the majority of currently recognized subspecies are not morphometrically distinct. In contrast, the separation between Bornean and Sumatran populations of T. tana is sufficient to recognize them as different subspecies. Comparisons of Bornean specimens to those on small, offshore islands reveal that the latter average smaller body size. This pattern is inconsistent with Foster’s island rule, which predicts that island populations of small mammals (< 5 kg) will average larger body size relative to mainland forms. A similar lack of support for ecogeographic rules has been noted in T. glis (Diard, 1820), suggesting that these “rules” are poor predictors of geographic variation in treeshrews.

Journal of Mammalogy

A new species of tree hyrax (Procaviidae: Dendrohyrax) from West Africa and the significance of the Niger–Volta interfluvium in mammalian biogeography

Tree hyraxes ( Dendrohyrax ) are one of only three genera currently recognized in Procaviidae, the only extant family in the mammalian order Hyracoidea. Their taxonomy and natural history have received little attention in recent decades. All tree hyrax populations of Guineo-Congolian forests of Africa are currently treated as a single species, Dendrohyrax dorsalis , the western tree hyrax, but many other groups of mammals distributed across this large biome have been shown to consist of several different species, each restricted to a distinct biogeographical region. We analysed variation in loud-call structure, pelage colour, skull morphometrics and mitochondrial genomes in populations across much of the range of D. dorsalis . This integrative approach uncovered considerable cryptic variation. The population found between the Niger and Volta Rivers in West Africa is particularly distinctive, and we describe it herein as a new species. Our study highlights the need to revise the taxonomy of the genus Dendrohyrax in light of modern systematics and current understanding of its distribution. It also adds to a growing body of evidence that the Niger–Volta interfluvium has a distinct meso-mammal fauna. Unfortunately, the fauna of this region is under major threat and warrants much greater conservation attention.

Zoological Journal of the Linnean Society

The limitations of external measurements for aging small mammals: The cautionary example of the Lesser Treeshrew (Scandentia: Tupaiidae: Tupaia minor Günther, 1876)

Age is a basic demographic characteristic vital to studies of mammalian social organization, population dynamics, and behavior. To eliminate potentially confounding ontogenetic variation, morphological comparisons among populations of mammals typically are limited to mature individuals (i.e., those assumed to have ceased most somatic growth). In our morphometric studies of treeshrews (Scandentia), adult individuals are defined by the presence of fully erupted permanent dentition, a common criterion in specimen-based mammalogy. In a number of cases, however, we have had poorly sampled populations of interest in which there were potentially useful specimens that could not be included in samples because they lacked associated skulls. Such specimens typically are associated with external body and weight measurements recorded by the original collectors, and we sought to determine whether these data could be used successfully as a proxy for age or at least to establish maturity. We analyzed four traditional external dimensions (head-and-body length, tail length, hind foot length, and ear length) and weight associated with 103 specimens from two allopatric populations of the Lesser Treeshrew ( Tupaia minor Günther, 1876) from Peninsular Malaysia and from Borneo, which we treated as separate samples (populations). Individuals were assigned to one of eight age categories based on dental eruption stage, and measurements were compared among groups. In general, mean sizes of infants and subadults were smaller than those of adults, but the majority of subadults fell within the range of variation of adults. The large overlap among infants, subadults, and adults in external measurements and weight indicates that such measures are poor proxies for age in this species, probably for treeshrews in general, and possibly for other small mammals. This has significant implications for any investigation wherein relative age of individuals in a given population is an important consideration.

Journal of Mammalogy

A new addition to the embalmed fauna of ancient Egypt: Güldenstaedt’s White-toothed Shrew, Crocidura gueldenstaedtii (Pallas, 1811) (Mammalia: Eulipotyphla: Soricidae)

The Falcon Necropolis at Quesna in the Nile Delta of Egypt is considered to have been founded by the priest Djedhor, the Saviour, of Athribis (Tell Atrib in modern Benha) at the beginning of the Ptolemaic Period. Recent excavations here have revealed abundant avian remains from mummies dedicated to the ancient Egyptian god Horus Khenty-Khety. Among the few mammal remains from the site are five species of shrews (Eulipotyphla: Soricidae), including some that we identified as Güldenstaedt’s White-toothed Shrew, Crocidura gueldenstaedtii (Pallas, 1811). Discovery of this species at Quesna increases the number of shrews recovered from ancient Egyptian archaeological sites to eight species. Crocidura gueldenstaedtii no longer occurs in the Nile Delta, and its presence in a diverse shrew fauna at Quesna that includes one other extirpated species, Crocidura fulvastra (Sundevall, 1843), supports the hypothesis of a moister regional environment 2000–3000 years ago. Inadvertently preserved local faunas, such as that from Quesna, can provide valuable information about ancient environments and subsequent turnover in faunal communities.

PLoS ONE

The green mole, Astromycter prasinatus T. M. Harris, 1825 (Mammalia: Eulipotyphla: Talpidae): An origin story

Thaddeus William Harris described the green mole of Maine, Condylura prasinata (Mammalia: Eulipotyphla: Talpidae), in the July 1825 issue of the Boston Journal of Philosophy and the Arts , and this was considered the original description of the species throughout most of the nineteenth century. In the early twentieth century, taxonomists began instead to cite an earlier notice in the June 1825 issue of the American Journal of Science and Arts . This short article also described the species, but also established a separate genus for it, Astromycter , despite Harris’s indications elsewhere that the species was congeneric with the star-nosed mole, Condylura cristata (Linnaeus, 1758). Moreover, the American Journal of Science and Arts article cited as its source the “ Machias Star ”, indicating the possibility of an even earlier description of the animal. With Astromycter prasinata in synonymy with C. cristata for over a century, little effort has been exerted to determine whether earlier source materials exist or why, within two months, the green mole was allocated to two different genera by its describer. The question is taxonomically relevant today because C. prasinata predates C. cristata nigra ( Smith, 1940), as an available name for north-eastern populations of star-nosed moles. If subspecies of C. cristata are to be recognized, the north-eastern subspecies should correctly bear the name C. cristata prasinata . In fact, authority for both genus- and species-group names for the green mole have been misattributed since 1825. The descriptions of C. prasinata by Thaddeus William Harris in the Boston Journal of Philosophy and the Arts and American Journal of Science and Arts were preceded by at least three published descriptions of A. prasinatus by Thaddeus Mason Harris, his father.

Archives of Natural History

Ancient Egyptian mummified shrews (Mammalia: Eulipotyphla: Soricidae) and mice (Rodentia: Muridae) from the Spanish Mission to Dra Abu el-Naga, and their implications for environmental change in the Nile valley during the past two millennia

Excavation of Ptolemaic Period (ca. 309–30 BC) strata within Theban Tombs 11, 12, -399-, and UE194A by the Spanish Mission to Dra Abu el-Naga (also known as the Djehuty Project), on the west bank of the Nile River opposite Luxor, Egypt, yielded remains of at least 175 individual small mammals that include four species of shrews (Eulipotypha: Soricidae) and two species of rodents (Rodentia: Muridae). Two of the shrews ( Crocidura fulvastra and Crocidura pasha ) no longer occur in Egypt, and one species ( Crocidura olivieri ) is known in the country only from a disjunct population inhabiting the Nile delta and the Fayum. Although deposited in the tombs by humans as part of religious ceremonies, these animals probably derived originally from local wild populations. The coexistence of this diverse array of shrew species as part of the mammal community near Luxor indicates greater availability of moist floodplain habitats than occur there at present. These were probably made possible by a greater flow of the Nile, as indicated by geomorphological and palynological evidence. The mammal fauna recovered by the Spanish Mission provides a unique snapshot of the native Ptolemaic community during this time period, and it permits us to gauge community turnover in the Nile valley of Upper Egypt during the last 2000 years. It also serves as a relevant example for understanding the extinction and extirpation of mammal species as effects of future environmental changes predicted by current climatic models.

northern Egypt

Coming of age: Morphometric variation in the hand skeletons of juvenile and adult Lesser Treeshrews (Scandentia: Tupaiidae: Tupaia minor Günther, 1876)

Morphometric analyses of the manus skeleton have proven useful in understanding species limits and morphological divergence among tupaiid treeshrews (Scandentia: Tupaiidae). Specimens in these studies are typically limited to mature individuals with fully erupted permanent dentition, which eliminates potentially confounding variation attributable to age, but can also exclude rare taxa and small island populations that are poorly represented in systematic collections. To determine the real limits associated with including immature animals in such studies, we used multivariate analyses to study sexual and age variation of the manus skeleton within two allopatric populations of the Lesser Treeshrew (Tupaia minor Günther, 1876) from the Malay Peninsula and from Borneo that we treated as separate samples. Individuals were aged using dental eruption of the permanent dentition. We also recorded the degree of epiphyseal fusion of the bones of the manus based on x-rays of study skins. We then tested our ability to distinguish the two populations using a series of discriminant function analyses of hand measurements from samples that included varying proportions of immature individuals and adults. We found no evidence of sexual dimorphism in hand proportions, permitting us to combine females and males in our samples. Epiphyseal fusion of the metacarpals and phalanges typically occurs by the time the third molars have completely erupted, and fusion of the distal epiphyses of the radius and ulna typically occurs by the time the permanent fourth premolars are in place. There is occasional asynchrony between dental age and epiphyseal fusion. In both populations, the hands of most infants and subadults provide morphometric values within the range of variation of adults, although they are typically distributed in the lower part of the adult range and have the potential to bias the sample toward lower mean size. The inclusion of infants and subadults when attempting to discriminate between two taxa generally results in lower rates of correct classifications, although the rates increase as the sample of immature individuals is limited to older subadults. As a general rule, we recommend that infants and subadults continue to be excluded from analyses when exploring taxonomic boundaries among treeshrews. In cases of extremely small sample sizes of adults, however, older subadults—in which the permanent third premolars are erupting or in place—can be used with appropriate caution.

Journal of Mammalogy

“Mostri Marini”: Constantine S. Rafinesque's names for three of Antonino Mongitore's Sicilian whales

In 1815, the naturalist Constantine Samuel Rafinesque (1783–1840) previewed three new species of cetaceans – Delphinus dalippus , Physeter urganantus and Oxypterus mongitori – that he intended to describe from Sicily based on illustrations in Antonino Mongitore's published work Della Sicilia ricercata nelle cose più memorabili (1742–1743). Although formal descriptions of the three species were never published, Rafinesque's reference to Mongitore's illustrations made the names available by “indication”. The names, nonetheless, fell into obscurity, most likely a result of contemporary taxonomists' lack of access to Mongitore's work. Rafinesque's names remain relevant to the history of cetacean taxonomy, although they are no longer applicable. Moreover, the animals associated with these names add to the historical record of whale strandings in the Mediterranean. For these reasons, we studied the illustrations Rafinesque indicated for his cetaceans and reviewed Mongitore's accompanying text, which together provide sufficient distinctive characters that two of the three animals can be confidently identified with modern species, namely the sperm whale, Physeter catodon ( Linnaeus, 1758), and the false killer whale, Pseudorca crassidens (Owen, 1846). Had Rafinesque's name D. dalippus been recognized for what it was, it would have had priority over P. crassidens as the earliest scientific name for the false killer whale.

Archives of Natural History

Comparative functional skeletal morphology among three genera of shrews: Implications for the evolution of locomotor behavior in the Soricinae (Eulipotyphla: Soricidae)

The clade comprising the soricid tribes Blarinellini ( Blarinella ) and Blarinini ( Blarina and Cryptotis ) is notable within the Soricidae (Eulipotyphla) for the large proportion of reportedly semifossorial species. To better define locomotor modes among species in these two tribes, we quantified purported locomotor adaptations by calculating 23 functional indices from postcranial measurements obtained from museum specimens of Blarina and Blarinella and published measurements for 16 species of Cryptotis . We then analyzed relative ambulatory–fossorial function of each species using principal component analyses and mean percentile rank (MPR) analysis of the indices. Species within the Blarinellini–Blarinini clade exhibit a graded series of morphologies with four primary functional groupings that we classified as “ambulatory,” “intermediate,” “semifossorial,” and “fossorial.” To obtain a preliminary overview of evolution of locomotor modes in this group, we mapped MPRs on a composite phylogeny and examined the resulting patterns. That analysis revealed that the most recent common ancestor of the Blarinellini–Blarinini clade most likely had an intermediate or semifossorial locomotor morphology. Individual subclades subsequently evolved either more ambulatory or more fossorial morphologies. Hence, evolution of locomotor traits within this clade is complex. Multiple shifts in locomotor mode likely occurred, and no single directional tendency is apparent either among the major modes or in levels of complexity.

Journal of Mammalogy

See how they ran: Morphological and functional aspects of skeletons from ancient Egyptian shrew mummies (Eulipotyphla: Soricidae: Crocidurinae)

Animals served important roles in the religious cults that proliferated during the Late (ca. 747–332 BCE) and Greco-Roman periods (332 BCE–CE 337) of ancient Egypt. One result was the interment of animal mummies in specialized necropolises distributed throughout the country. Excavation of a rock-tomb that was re-used during the Ptolemaic Period (ca. 309–30 BCE) for the interment of animal mummies at the Djehuty Site (TT 11-12) near Luxor, Egypt, was carried out in early 2018 by a Spanish-Egyptian team sponsored by the Consejo Superior de Investigaciones Científicas, Madrid. The tomb burned sometime after deposition of the mummies, leaving behind abundant disassociated skeletal remains, primarily of avians, but also including two species of shrews (Soricidae): Crocidura olivieri (Lesson, 1827) and C. religiosa (I. Geoffroy Saint-Hilaire, 1826). To investigate possible intraspecific variation in morphology and locomotor function in these two species during the last two millennia, we measured morphological features of individual postcranial bones from the two archaeological samples and calculated indices that have been used to assess locomotor function. We compared the measurements to those from modern C. olivieri , C. religiosa , and C. suaveolens (Pallas, 1811) using principal components analysis, and we compared locomotor indices to those we calculated for the three modern species of Crocidura and to those from nine species of myosoricine shrews. Osteological features of the postcranial skeleton of conspecific Ptolemaic and modern samples of C. olivieri and C. religiosa are generally similar in character and proportion, and, skeletally, these shrews and modern C. suaveolens are consistent with soricids having a primarily ambulatory locomotor mode. One exception is the deltopectoral crest of the humerus, which appears to be longer in modern C. religiosa. Despite general conservation of form and function, Ptolemaic C. olivieri had larger body size than modern Egyptian populations and were more similar in size to modern C. olivieri nyansae , 1900 from Kenya than to modern C. olivieri olivieri from Egypt.

Luxor

Shrews (Eulipotyphla, Soricidae) of Guatemala /Musarañas (Eulipotyphla, Soricidae) de Guatemala

Shrews (Soricidae) are the only members of the mammalian order Eulipotyphla that occur in Central and South America. In Guatemala, 15 species have been recorded belonging to the genera Cryptotis and Sorex , three of which are new and undescribed. Two additional species are expected to be discovered in the country based on their known distributions. Most species appear to have limited reproduction throughout the year. We review the taxonomy and natural history of these species.

Book chapter