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.

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.

Mitogenome and phylogenetic analyses support rapid diversification among species groups of small-eared shrews genus Cryptotis (Mammalia: Eulipotyphla: Soricidae).

The small-eared shrew genus Cryptotis is the third largest in the family Soricidae and occurs in North, Central, and northern South America. In Mexico and Central and South America, most species inhabit geographically isolated moist, montane habitats at middle and high elevations in a typical sky-island pattern. The 49 recognized species have been partitioned into as many as six species groups based on morphological and molecular phylogenetic studies. The relationships among these species groups are poorly resolved, and their evolutionary histories, including migration patterns and locomotor adaptations, remain unclear. Herein, we provide a new phylogeny incorporating complete mitochondrial genomes (mitogenomes) and supermatrix approach. We compared different evolutionary scenarios using approximately unbiased (AU), Kishino-Hasegawa (KH), and Shimodaira-Hasegawa (SH) statistical tests. The phylogenetic hypothesis based on mitogenomes revealed novel relationships supporting a basal position for the Cryptotis parvus-group in the genus, and a close relationship between C. gracilis and one clade of the C. thomasi-group. The former relationship is consistent with the least derived humerus morphology and northern distribution of the species. The latter relationship implies multiple migrations between Central and South America. The lack of fine resolution for the species group relationships may be due partly to the lack of taxon sampling. In contrast, multi-approach analyses suggest that the unresolved relationships may be a result of rapid diversification during the early stages of Cryptotis evolution.

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.

Rule reversal: Ecogeographical patterns of body size variation in the common treeshrew (Mammalia, Scandentia).

There are a number of ecogeographical "rules" that describe patterns of geographical variation among organisms. The island rule predicts that populations of larger mammals on islands evolve smaller mean body size than their mainland counterparts, whereas smaller-bodied mammals evolve larger size. Bergmann's rule predicts that populations of a species in colder climates (generally at higher latitudes) have larger mean body sizes than conspecifics in warmer climates (at lower latitudes). These two rules are rarely tested together and neither has been rigorously tested in treeshrews, a clade of small-bodied mammals in their own order (Scandentia) broadly distributed in mainland Southeast Asia and on islands throughout much of the Sunda Shelf. The common treeshrew, Tupaia glis, is an excellent candidate for study and was used to test these two rules simultaneously for the first time in treeshrews. This species is distributed on the Malay Peninsula and several offshore islands east, west, and south of the mainland. Using craniodental dimensions as a proxy for body size, we investigated how island size, distance from the mainland, and maximum sea depth between the mainland and the islands relate to body size of 13 insular T. glis populations while also controlling for latitude and correlation among variables. We found a strong negative effect of latitude on body size in the common treeshrew, indicating the inverse of Bergmann's rule. We did not detect any overall difference in body size between the island and mainland populations. However, there was an effect of island area and maximum sea depth on body size among island populations. Although there is a strong latitudinal effect on body size, neither Bergmann's rule nor the island rule applies to the common treeshrew. The results of our analyses demonstrate the necessity of assessing multiple variables simultaneously in studies of ecogeographical rules.

Rediscovery of the type series of the Sacred Shrew, Sorex religiosus I. Geoffroy Saint-Hilaire, 1826, with additional notes on mummified shrews of ancient Egypt (Mammalia: Soricidae).

In 1826, Isidore Geoffroy Saint-Hilaire described the Sacred Shrew, Sorex religiosus [= Crocidura religiosa] from a series of 22 embalmed individuals that comprised a portion of the Italian archeologist Joseph Passalacqua's collection of Egyptian antiquities from an ancient necropolis near Thebes, central Egypt. Living members of the species were not discovered until the beginning of the 20th century and are currently restricted to the Nile Delta region, well north of the type locality. In 1968, the type series of S. religiosus was reported lost, and in 1978, a neotype was designated from among a small collection of modern specimens in the Natural History Museum, London. Our investigations have revealed, however, that the type series is still extant. Most of the specimens used by I. Geoffroy Saint-Hilaire to describe S. religiosus still form part of the Passalacqua Collection in the Ägyptisches Museum, Berlin, Germany. We summarize the taxonomic history of S. religiosus, review the history of the Passalacqua collection, and explain why the type series was thought to have been lost. We designate an appropriate lectotype from among the original syntypes of S. religiosus in the Ägyptisches Museum. Our examination of the shrew mummies in the Passalacqua collection also yielded a species previously unrecorded from either ancient or modern Egypt: Crocidura pasha Dollman, 1915. Its presence increases the number of soricid species embalmed in ancient Egypt to seven and provides additional evidence for a more diverse Egyptian shrew fauna in the archeological past. Finally, we provide details that will assist in better understanding the variety of mummification procedures used to preserve animals in ancient Egypt.

Molecular Phylogeny Supports Repeated Adaptation to Burrowing within Small-Eared Shrews Genus of Cryptotis (Eulipotyphla, Soricidae).

Small-eared shrews of the New World genus Cryptotis (Eulipotyphla, Soricidae) comprise at least 42 species that traditionally have been partitioned among four or more species groups based on morphological characters. The Cryptotis mexicana species group is of particular interest, because its member species inhibit a subtly graded series of forelimb adaptations that appear to correspond to locomotory behaviors that range from more ambulatory to more fossorial. Unfortunately, the evolutionary relationships both among species in the C. mexicana group and among the species groups remain unclear. To better understand the phylogeny of this group of shrews, we sequenced two mitochondrial and two nuclear genes. To help interpret the pattern and direction of morphological changes, we also generated a matrix of morphological characters focused on the evolutionarily plastic humerus. We found significant discordant between the resulting molecular and morphological trees, suggesting considerable convergence in the evolution of the humerus. Our results indicate that adaptations for increased burrowing ability evolved repeatedly within the genus Cryptotis.

Functional skeletal morphology and its implications for locomotory behavior among three genera of myosoricine shrews (Mammalia: Eulipotyphla: Soricidae).

Myosoricinae is a small clade of shrews (Mammalia, Eulipotyphla, Soricidae) that is currently restricted to the African continent. Individual species have limited distributions that are often associated with higher elevations. Although the majority of species in the subfamily are considered ambulatory in their locomotory behavior, species of the myosoricine genus Surdisorex are known to be semifossorial. To better characterize variation in locomotory behaviors among myosoricines, we calculated 32 morphological indices from skeletal measurements from nine species representing all three genera that comprise the subfamily (i.e., Congosorex, Myosorex, Surdisorex) and compared them to indices calculated for two species with well-documented locomotory behaviors: the ambulatory talpid Uropsilus soricipes and the semifossorial talpid Neurotrichus gibbsii. We summarized the 22 most complete morphological variables by 1) calculating a mean percentile rank for each species and 2) using the first principal component from principal component analysis of the indices. The two methods yielded similar results and indicate grades of adaptations reflecting a range of potential locomotory behaviors from ambulatory to semifossorial that exceeds the range represented by the two talpids. Morphological variation reflecting grades of increased semifossoriality among myosoricine shrews is similar in many respects to that seen for soricines, but some features are unique to the Myosoricinae.

Can they dig it? Functional morphology and semifossoriality among small-eared shrews, genus Cryptotis (Mammalia, Soricidae).

Small-eared shrews (Mammalia: Soricidae: Cryptotis), exhibit modifications of the forelimb skeleton that have been interpreted as adaptations for semifossoriality. Most species inhabit remote regions, however, and their locomotory and foraging behaviors remain mostly speculative. To better understand the morphological modifications in the absence of direct observations, we quantified variation in these species by measuring 151 individuals representing 18 species and populations of Cryptotis and two species of moles (Talpidae) for comparison. From our measurements, we calculated 22 indices, most of which have been used previously to characterize substrate use among rodents and other taxa. We analyzed the indices using 1) average percentile ranks, 2) principal components analysis, and 3) cluster analysis. From these analyses, we determined that three basic modes of substrate adaptation are present within Cryptotis: 1) a primarily terrestrial mode, with species that are capable of burrowing, but lack adaptations to increase digging efficiency, 2) a semifossorial mode, with species whose forelimbs bones show strong muscle attachment areas and increased mechanical advantage, and 3) an intermediate mode. In addition to identifying new morphological characters and contributing to our understanding of the functional morphology of soricids, these analyses provide additional insight into the ecology of the species of interest.

Survival of the less-fit: a least shrew (Mammalia, Soricidae, Cryptotis parvus) survives a separated leg fracture in the wild.

X-rays of the deformed femur of a least shrew (Cryptotis parvus) revealed an ossified callus enclosing a greatly displaced, spiral fracture. Based on models of fracture healing, this shrew lived for at least 28 days postfracture. Survival of major fractures in the wild may not be uncommon.

Skeletal morphology of the forefoot in shrews (Mammalia: Soricidae) of the genus Cryptotis, as revealed by digital X-rays.

Variation in the forefoot skeleton of small-eared shrews (family Soricidae, genus Cryptotis) has been previously documented, but the paucity of available skeletons for most taxa makes assessment of the degrees of intraspecific and interspecific variation difficult. We used a digital X-ray system to extract images of the forefoot skeleton from 101 dried skins of eight taxa (seven species, including two subspecies of one species) of these shrews. Lengths and widths of each of the four bones of digit III were measured directly from the digital images, and we used these data to quantify variation within and among taxa. Analysis of the images and measurements showed that interspecific variation exceeds intraspecific variation. In fact, most taxa could be distinguished in multivariate and some bivariate plots. Our quantitative data helped us define a number of specific forefoot characters that we subsequently used to hypothesize evolutionary relationships among the taxa using the exhaustive search option in PAUP, a computer program for phylogenetic analysis. The resulting trees generally concur with previously published evolutionary hypotheses for small-eared shrews. Cryptotis meridensis, a taxon not previously examined in recent phylogenies, is rooted at the base of the branch leading to the C. mexicana group of species. The position of this species suggests that the mostly South American C. thomasi group shares an early ancestor with the C. mexicana group.