Macrostructure and evolution of the digestive system in Echinoida (Echinodermata)

The structure of the digestive system in Echinoida has long been puzzling since comparative studies have suggested that a derived structure, the siphon, has apparently evolved twice independently. New observations on the digestive system in five species of Cidaroida, four species of the Diadematoida and three species of Echinothurioida are presented. The results show that the four diadematoid species have a siphon and the three species of Echinothurioida have a siphonal groove, contrary to previous assertions. These observations make the macrostructure of the echinoid digestive system fully consistent with more recent phylogenetic hypotheses based on molecular and general morphological data, and support the idea that a siphon has evolved only once, in the stem lineage of the Acroechinoidea.     

Recovery from Siphon Damage in Donax vittatus (Da Costa) (Bivalvia: Donacidae)

Between 2.5% and 18% of Donax vittatus from a natural populationon West Sands beach, St Andrews on the Scottish east coast showeddamage to the siphons caused by non-lethal predation by juvenileflatfishes. The percentage with damaged siphons was greatestin summer. Either inhalant or exhalant siphons were affected,in varying proportions, or in some cases, both siphons. In experimentalaquaria, Donax vittatus that suffered non-lethal attack by juvenileplaice which resulted in their removal from the sand, rapidlyresumed normal activity as evidenced by reburrowing. Wound healing,followed by re-differentiation of siphonal tentacles, took placerapidly following experimental amputation of siphon tips, withthe newly-formed tentacles appearing almost normal after 10days. Re-differentiation of the siphonal tentacles was accompaniedby the development of their complement of three types of ciliated senseorgans. (Received 11 May 1998; accepted 9 July 1998)     

As good as dead? Sublethal predation facilitates lethal predation on an intertidal clam

Although ecologists have speculated that sublethal predation can impact prey dynamics, consequences of these predator effects have seldom been experimentally tested. In soft‐sediment marine communities, fishes crop extended feeding siphons of buried clams, potentially causing clams to reduce their burial depth, thereby enhancing their susceptibility to excavating lethal predators. We simulated cropping of the confamilial clams, Protothaca staminea and Venerupis philippinarum, by removing the top 40% of siphons, which caused each species to burrow 33–50% shallower than conspecifics with intact siphons. To examine subsequent consequences of reduced burial depth, we exposed cropped and intact clams to natural levels of predation in the field. Because of a naturally longer siphon, Protothaca, even after cropping, remained at relatively safe burial depths. In contrast, siphon cropping nearly doubled the mortality rate of Venerupis. Thus, while sublethal predation facilitates lethal predation, this linkage depends on specific life history characteristics, even among ecologically similar species.     

SOME BEHAVIOURAL AND ELECTRICAL RESPONSES OF SCROBICULARIA PLANA (BIVALVIA:TELLINACEA) TO SIPHONAL WOUNDING

Water pumping, valve movements and heart rate have been recordedfrom Scrobicularia for short periods of normal behaviour andthen after siphonal wounding. Scrobicularia exhibits regularand repetitive pumping periods interrupted for only 2–3s after siphonal wounding, without the regularity of these periodsbeing affected. Wounding does not prevent animals from usingtheir inhalant siphons for deposit feeding. A preliminary investigationof neural responses to stimulation has shown that wounding thesiphon causes minimal disturbance to the animal, a brief (2s)burst of nerve activity occurs, the siphon is retracted, butvalve adduction does not occur. In contrast to this tactilestimulation of the mantle edge always elicits a large burstof impulses in the posterior adductor nerve, valve closure results,usually for 14–15 s. ¹Present address: Dept of Zoology, University of Cape Town,Rondebosch 7700, South Africa. (Received 2 February 1981;     

A revision of some Asian and African freshwater clams assigned to Type=”Italic”>Corbicula fluminalis (Müller, 1774) (Mollusca: Bivalvia: Corbiculidae), with a review of anatomical characters and reproductive features based on museum collections

A study of museum specimens of the genus Corbicula from the western Asiatic part of its range (Middle East, the Caucasus and Central Asia) has provided the following diagnostic anatomical characters for Corbicula fluminalis (Müller, 1774): siphons narrow with circular apertures; siphonal papillae rather scarce: about 40 around inhalant siphon (one or two rows) and 12--20 around exhalant siphon (one incomplete row); ring of dark pigment usually present internally at base of each siphon; papillae of fused mantle lobes and free mantle edge numerous, the first organised in one or two rows; radial muscles of mantle edge well developed and arranged in bands. Presence of hermaphroditic specimens, biflagellate type of spermatozoa indicating clonality, and incubation of D-shaped 190--217 m long larvae in gills were also shown for this taxon. Two conchologically different morphotypes referred to in literature as C. fluminalis s. str. and C. purpurea (Prime, 1864) show also slight differences in size of spermatozoa and may represent different clonal lineages. The lots from North Africa possess the same characters as one of the Asian morphotypes, thus taxonomic distinctness of C. fluminalis consobrina (Cailliaud, 1827) is not supported. In contrast, C. africana (Krauss, 1848) distributed in Lake Malawi and South Africa and treated in some modern reviews as another subspecies of C. fluminalis differs by the patterns of shell sculpture, the lower number of siphonal papillae, the absence of pigmentation on the siphons and their papillae, and the size of spermatozoa; its distinctness is therefore supported, but close affinity to C. fluminalis needs confirmation. All studied African morphotypes are identical in their mode of reproduction. The new data on the anatomy and reproduction of C. fluminalis show strong similarities between this taxon and the eastern incubating species C. fluminea (Müller, 1774), but these taxa can be distinguished by their shell characters. The Chinese estuarine non-incubating Corbicula earlier assigned to C. fluminalis shows different anatomical and reproductive characters and is re-considered as C.cf. japonica (Prime, 1864). These results are discussed in the framework of Corbicula phylogeny based on molecular characters.     

Control of foraging behavior of individuals within an ecosystem context: the clam Macoma balthica,flow environment,and siphon-cropping fishes

Macoma balthica (L.), an abundant clam, ubiquitous in temperate estuaries across the North Atlantic, is known to practice both alternative basic modes of feeding available to seafloor invertebrates. It either holds its feeding organ, the siphon, at a fixed position just above the sediment surface to filter out food particles suspended in the overlying water or else extends and moves its siphon around to vacuum up deposited food particles on the sediment surface. Previous laboratory experiments have established an understanding of the role of current flow in dictating the choice of whether suspension or deposit feeding will be used by marine invertebrates with the facultative flexibility to choose. Faster flows imply greater fluxes of suspended particles so that the energetic rewards of suspension feeding are enhanced. Slower flows imply reduced renewal rates of suspended foods in the bottom boundary layers and enhanced deposition of food particles on the seafloor so that a switch to deposit feeding is favored. Like early optimal foraging theory, this understanding is based on energetic considerations alone without incorporation of broader implications of how population interactions such as predation and competition influence individual foraging behavior. Feeding behavior of Macoma balthica is influenced in the Neuse River estuary by both hydrodynamics and siphon-cropping by juvenile demersal fishes. Under conditions of identical concentrations of suspended particulates in the water column and organic contents of surface sediments, Macoma exhibited much higher levels of deposit feeding where currents were slower. In addition, exclosure and fish inclosure experiments demonstrated that juvenile demersal fishes influence feeding behavior of Macoma by cropping exposed siphons and inducing reduction in deposit-feeding activity. Effects of croppers were substantial in early to midsummer, when juvenile fish abundances were greatest in trawl samples from this estuarine nursery and before the growing fish exhibited ontogenetic changes in diet away from early concentration on bivalve siphons. Field experiments in which siphon-cropping fish were caged at varying distances off the bottom failed to detect any effective behavioral avoidance by Macoma of cropping in response to proximity of fish. One might have hypothesized that under high risk of cropping, Macoma would switch to suspension feeding and away from deposit feeding, the feeding method entailing more risk of losses to croppers because of greater siphon activity and greater extension of siphons on the sediment surface. Consequently, partial predation by siphon-cropping fishes greatly reduces deposit-feeding activity by Macoma balthica during summer as an apparent direct effect of disfigurement and reduction of siphons, the organ required for efficient deposit feeding. Information on current flows alone would not suffice to predict feeding behavior of this marine invertebrate: the influence of partial predation must also be included.     

Arenophilic mantle glands in the Laternulidae (Bivalvia: Anomalodesmata) and their evolutionary significance

The mantle margins of several anomalodesmatans bear multicellular arenophilic glands, the mucoid secretions of which attach sand grains and other foreign particles to the outer surface of the periostracum. These glands have been recorded for many of the anomalodesmatan families and are used as a key morphological character in recent attempts to unravel the evolutionary relationships within the Anomalodesmata. The glands occur in Laternula elliptica, L. truncata, L. boschasina and L. marilina, discharging from the top of muscular papillae at the distal tip of the siphons. The secretions are laid down as threads organized in longitudinal lines along the length of the periostracum that covers the siphonal walls. This is the first record of arenophilic mantle glands in members of the Laternulidae, a finding that not only broadens our current knowledge of the family's morphology, but assists in the reconstruction of anomalodesmatan evolutionary history.     

Non-respiratory podia of clypeasteroids (Echinodermata,Echinoides): II. Diversity

Summary There are five major types of non-respiratory podia in the Order Clypeasteroida: accessory, barrel-tipped, food groove, large food groove, and buccal. The anatomy of each type is intimately related to its function in the feeding mechanism of clypeasteroids. Accessory podia are found aborally and orally in some species, only aborally and ambitally in others. Accessory podia are largely sensory and manipulatory, but can be locomotory in the small fibulariids and juvenile sand dollars. Barrel-tipped podia have expanded disk muscles and connective tissue, and are usually found in two sizes, large and small. In species that have them they are usually restricted to the oral surface. These podia collect food and pass it towards the food grooves in the manner of a bucket brigade. Food groove podia are found only in species with food grooves. These podia are small, with reduced tip musculature and expanded secretory tissue for coating food with mucus. They transport food down the food grooves to the mouth. Large food groove podia are simply large versions of ordinary food groove podia. They help move the clumped food into the mouth area towards the buccals, and are found only in the Clypeasteridae and some scutellines. Buccal podia lack tip musculature, but possess tip support fibres and a single type of small secretory cell. They are sensory, and capable of manipulating particles into the mouth. Buccals are present in all families except the Clypeasteridae. Juvenile Echinarachnius less than 3 mm in diameter have only respiratory, accessory and buccal podia. Food groove and barrel-tipped podia start to differentiate from the accessories as the juvenile approaches a diameter of 4 to 5 mm. Clypeasteroid podial diversity increases the efficiency of the food collecting mechanism. The anatomy and distribution of podia on the oral surface of scutellines supports the fact that this surface is the prime food collecting area in all true sand dollars. The podia (not miliary spines) are the major source of mucus used during the feeding process and are the primary feeding appendages.     

Regeneration of lost siphon tissues in the tellinacean bivalve Nuttallia olivacea

The inhalant siphon of the tellinacean bivalve Nuttallia olivacea is an important prey item for juvenile stone flounder Platichthys bicoloratus in estuaries in Japan. We examined quantitative siphon regeneration of N. olivacea in rearing experiments of siphon-removed bivalves (> 30 mm shell length) both in the laboratory and in their natural habitat. Under laboratory conditions, siphon-removed bivalves regenerated lost tissues quantitatively at 15 and 25 °C 1 mo after siphon removal, although regeneration was incomplete. A 3-mo caging experiment in the field showed that great regeneration occurred in siphon-removed bivalves. However, the siphon weight of removed bivalves was significantly smaller than that of non-amputated bivalves, suggesting the incomplete regeneration. In a 1-mo caging experiment, bivalves that had approximately 15% of their siphons amputated were selected at some intervals to illustrate the quantitative regeneration process. Estimated daily siphon production was remarkably high only a few days after amputation. It decreased greatly thereafter, but regeneration was not completed within 30 d. These results indicate that bivalves regenerate siphons rapidly just after losing siphon tissues and then regeneration is slowed down before it is completed.     

Protection against suspended sand: the function of the branchial membrane in the blue mussel Mytilus edulis

Blue mussels (Mytilus edulis) living in estuaries have to cope with varying concentrations of suspended sand. Sand flowing through the inhalant siphons comes into the infrabranchial chamber. The inhalant siphon can be partially closed by the branchial membrane. As a result the inward flow decreases, and suspended sand sinks and can be eliminated. Experiments with mussels from three ecologically different locations showed about the same response of the branchial membrane on contact with suspended sand. The presence and function of the branchial membrane appears to be an adaptation of mussels to their estuarine environment.     

Effects of green algal mats on bivalves in a New England mud flat

Concurrent with the spread of green algal mats on tidal flats, reports of macrofauna dieoffs under dense algal mats have increased in numbers. Bivalves seem to be particularly affected by persistent dense algal mats. Bivalve species with a long extendible siphon seem to be less affected underneath algal mats, but no distinction has been made in the past between species with short and those with long siphons,Mya arenaria andMacoma balthica, on an intertidal mudflat in New England. Abundances ofM. arenaria declined substantially during the study period when a thick green algal mat covered the mudflat for several months. Numbers of the small bivalveGemma gemma also decreased substantially, whereas abundances ofM. balthica showed minimal variation during the time of algal coverage. In algae removal/addition experiments numbers ofM. arenaria decreased, but effects were only significant in an algal addition to previously algal-free mudflat areas. Abundance ofM. balthica did not change significantly in the algal removal/additition experiments. Over the time period of the experiment (9 weeks),M. arenaria showed measurable size increase in uncovered mudflat areas, but not underneath algal mats. Similarly,M. balthica only increased in size in the uncovered mudflat area. From these results it is concluded thatM. balthica can survive time periods of dense algal coverage because it is able to penetrate through the algal mat with its long extendible siphon, and thus can reach well-oxygenated water layers above the mat.M. arenaria with its thick, less extendible, siphon cannot push through dense algal mats and therefore is more likely to die underneath persistent algal mats.     

Autocatalysis in Reaction Networks

The persistence conjecture is a long-standing open problem in chemical reaction network theory. It concerns the behavior of solutions to coupled ODE systems that arise from applying mass-action kinetics to a network of chemical reactions. The idea is that if all reactions are reversible in a weak sense, then no species can go extinct. A notion that has been found useful in thinking about persistence is that of “critical siphon.” We explore the combinatorics of critical siphons, with a view toward the persistence conjecture. We introduce the notions of “drainable” and “self-replicable” (or autocatalytic) siphons. We show that: Every minimal critical siphon is either drainable or self-replicable; reaction networks without drainable siphons are persistent; and nonautocatalytic weakly reversible networks are persistent. Our results clarify that the difficulties in proving the persistence conjecture are essentially due to competition between drainable and self-replicable siphons.     

The colonic groove or furrow: A comparative morphological study of six species of African mole‐rats (Rodentia,Bathyergidae)

Herbivorous mammals such as nutrias, guinea pigs, chinchillas, and mole‐rats have a longitudinal mucosal colonic groove (furrow) in their ascending colon, which is thought to play a role in the colonic separation mechanism (CSM). It is not known whether this groove is structurally modified to adapt to this function in mole‐rat species. The morphology of this groove was studied in 32 mol‐rats, four species, one of which consisted of three subspecies, endemic to southern Africa and two species found in eastern Africa. The macroscopic morphology of the groove was documented, and samples for histological examination were taken. The groove was wide at its origin at the cecocolic junction and was lined on either side by a row of papillae with the opposing papillae slightly offset in arrangement. The papillated groove gradually decreased in size toward the distal part of the ascending colon where it disappeared. This pattern was similar in all species except in Heterocephalus glaber, where the papillae were absent and the groove was lined by two longitudinal ridges. A histological examination of cross sections revealed that the mucosa covering the inner and outer walls of the groove was rich in mucous‐secreting goblet cells. The walls of the groove contained smooth muscle extending from the inner circular smooth muscle layer at the base to the tips of the papillae in all species examined as well as arteries, lymphatic vessels, and prominent sinusoid‐like veins. The groove could be demonstrated both macroscopically and histologically in three Bathyergus suillus fetuses of varying sizes. The sinusoid‐like veins present in all grooves, regardless of macroscopic shape, suggest that they have a role in the functioning of the groove. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Control of foraging behavior of individuals within an ecosystem context: the clam Macoma balthica and interactions between competition and siphon cropping

Macoma balthica (L.) is a common clam of the estuarine seafloor, belonging to an important group of invertebrates possessing the capacity to choose between the two fundamental modes of feeding available, using its siphon to inhale either suspended food particles from the water or food particles deposited on the sediment surface. Field experiments demonstrate that intraspecific competition, effects of other competing benthic invertebrates, and complex interactions between competition and partial predation (siphon cropping by fishes) modify the foraging behavior of Macoma. When protected by caging from siphon nipping by fishes, Macoma demonstrated greater siphon regeneration at lower density, indicating the importance of competition for limited resources. In the absence of siphon croppers, these same clams also exhibited more deposit feeding at the lower density either because of improved ability to deposit-feed with longer siphons or because deposited foods become more rapidly depleted than suspended foods on local spatial scales. Addition of siphon-nipping fishes caused greater reductions in siphon size of clams at lower density, presumably because the intensity of nipping per clam was greater where clam targets were fewer and because deposit feeding, which was more intense at lower densities, confers a greater risk of cropping from greater siphon extension and activity than characterize suspension feeding. Deposit feeding by Macoma was reduced in the presence of siphon croppers at both high and low density of clams, but the intensity of deposit-feeding activity at low density was substantially higher than predicted by additive effects of clam density and cropping. This suggests operation of a balancing strategy in Macoma whereby it is accepting greater risks of partial predation when rewards of greater food harvest are larger. The surprising failure to adopt a risk-averse stratery may be explained by the non-lethal nature of partial predation, which renders siphon loss an energetic penalty replacable through regeneration. The presence of a bed of suspension-feeding Rangia cuneata also altered foraging behavior of Macoma by inducing a switch to more intense deposit feeding, in response either to the documented near-bottom depletion of suspended foods or to likely enhanced biodeposition from feces and pseudofeces. The induction of greater deposit feeding by the presence of this competing suspension feeder led to greater siphon losses during exposure to croppers because Macoma was practicing more risky feeding behavior. This enhanced loss of siphon tissues to croppers in the presence of the suspension-feeding Rangia induced an interaction between the effects of siphon croppers and Rangia, such that Macoma exhibited a larger switch away from deposit feeding in the presence of siphon croppers when Rangia were also present. Clearly, the foraging decisions made by individuals can only be understood in a broad holistic context of population, community, and ecosystem processes.     

Changes to the geometry and fluid mechanics of the carotid siphon in the pediatric Moyamoya disease

Background: The Moyamoya disease is a cerebrovascular disease that causes occlusion of the distal end of the internal carotid artery, leading to the formation of multiple tiny collateral arteries. To date, the pathogenesis of Moyamoya is unknown. Improved understanding of the changes to vascular geometry and fluid mechanics of the carotid siphon during disease may improve understanding of the pathogenesis, prognosis techniques and disease management. Methods: A retrospective analysis of Magnetic Resonance Angiography (MRA) images was performed for Moyamoya pediatric patients (MMD) (n = 23) and control (Ctrl) pediatric patients (n = 20). The Ctrl group was composed of patients who complained of headache and had normal MRA. We performed segmentation of MRA images to quantify geometric parameters of the artery. Computational fluid dynamics (CFD) was performed to quantify the hemodynamic parameters. Results: MMD internal carotid and carotid siphons were smaller in cross-sectional areas, and shorter in curved vascular length. Vascular curvature remained constant over age and vascular size and did not change between Ctrl and MMD, but MMD carotid siphon had lower tortuosity in the posterior bend, and higher torsion in the anterior bend. Wall shear stress and secondary flows were significantly lower in MMD, but the ratio of secondary flow kinetic energy to primary flow kinetic energy were similar between MMD and Ctrl. Conclusion: There were alterations to both the geometry and the flow mechanics of the carotid siphons of Moyamoya patients but it is unclear whether hemodynamics is the cause or the effect of morphological changes observed.     

Three new species of <Emphasis Type="Italic">Anthocephalum</Emphasis> Linton, 1890 (Cestoda: Tetraphyllidea) from dasyatid stingrays of the Gulf of California

Three new species of Anthocephalum Linton, 1890 are described from dasyatid stingrays collected in the Gulf of California. Anthocephalum michaeli n. sp. is described from Dasyatis longus (Garman). This species most closely resembles A. alicae Ruhnke, 1994, but differs from this species in proglottid number. A. lukei n. sp. is also described from D. longus. This new species is most similar to A. cairae Ruhnke, 1994, but differs from that species in marginal loculi number and number of proglottids. The third new species, A. currani n. sp., is described from D. brevis (Garman). This species is most similar to A. centrurum (Southwell, 1925) Ruhnke, 1994, but differs from that species in marginal loculi number, number of testes and ovarian length. Phyllobothrium kingae Schmidt, 1978 is also consistent in morphology with species of Anthocephalum and is transferred to this genus, forming the new combination Anthocephalum kingae n. comb. This species most closely resembles A. michaeli n. sp., but differs in testicular shape. This brings the total number of species of Anthocephalum to nine. The transfer of the species Phyllobothrium arctowskii Wojciechowska, 1991, P. georgiense Wojciechowska, 1991, P. rakusai Wojciechowska, 1991 and P. siedleckii Wojciechowska, 1991 to Anthocephalum is not warranted, as these four species lack a posteriorly recurved cirrus-sac and a sinuous vagina, and have vitelline follicles uninterrupted by the ovary. Of the nine known species, all are parasitic in batoid fishes, and six are found in species of Dasyatis Garman. The phylogenetic status of Anthocephalum species in relationship to Rhinebothroides Mayes, Brooks & Thorson, 1981, Pararhineothroides Zamparo, Brooks & Barriga, 1999 and other rhinebothriin taxa is discussed.     

Validity of <Emphasis Type="Italic">Scolecenchelys aoki</Emphasis>, with a redescription of <Emphasis Type="Italic">Scolecenchelys gymnota</Emphasis> (Anguilliformes: Ophichthidae)

The myrophine ophichthid fishes (worm eels) Muraenichthys aoki Jordan and Snyder 1901 and Muraenichthys gymnotus Bleeker 1857 are redescribed as valid species of Scolecenchelys based on the types and non-type specimens collected from the Indo-Pacific. Because both species are similar to each other in having acute snouts, the posterior margin of the eye before the rictus, and their dorsal-fin origins located slightly posterior to a vertical line through the anus, Scolecenchelys aoki has usually been regarded as a junior synonym of Scolecenchelys gymnota. However, S. aoki is clearly distinguishable from S. gymnota by having a median groove on the ventral side of snout (absent in S. gymnota), uniserial maxillary teeth in smaller specimens (<200 mm TL; vs. biserial), three infraorbital sensory pores at postorbital area (vs. two), and more numerous vertebrae (56–65 in predorsal vs. 51–57; 53–58 in preanal vs. 47–52). Scolecenchelys aoki is restricted to Japanese waters and regarded as a senior synonym of Muraenichthys borealis Machida and Shiogaki 1990. Scolecenchelys gymnota is widely distributed in the Indo-Pacific, from South Africa and the Red Sea to Samoa, north to Okinawa, Japan. Sphagebranchus huysmani Weber 1913 and Muraenichthys fowleri Schultz 1943 are synonymized under S. gymnota.     

Compartmental neural simulations with spatial adaptivity

Since their inception, computational models have become increasingly complex and useful counterparts to laboratory experiments within the field of neuroscience. Today several software programs exist to solve the underlying mathematical system of equations, but such programs typically solve these equations in all parts of a cell (or network of cells) simultaneously, regardless of whether or not all of the cell is active. This approach can be inefficient if only part of the cell is active and many simulations must be performed. We have previously developed a numerical method that provides a framework for spatial adaptivity by making the computations local to individual branches rather than entire cells (Rempe and Chopp, SIAM Journal on Scientific Computing, 28: 2139–2161, 2006). Once the computation is reduced to the level of branches instead of cells, spatial adaptivity is straightforward: the active regions of the cell are detected and computational effort is focused there, while saving computations in other regions of the cell that are at or near rest. Here we apply the adaptive method to four realistic neuronal simulation scenarios and demonstrate its improved efficiency over non-adaptive methods. We find that the computational cost of the method scales with the amount of activity present in the simulation, rather than the physical size of the system being simulated. For certain problems spatial adaptivity reduces the computation time by up to 80%.     

Organization of the vomeronasal organ in a plethodontid salamander

Salamanders in the family Plethodontidae show a unique behavior (nose-tapping) and have unique structures (nasolabial grooves) that may be used specifically to convey chemicals to the vomeronasal organ. The nasal structure of Plethodon cinereus was studied to determine if there is enhanced development of the vomeronasal organ compared with other salamander families that would correlate with use of these unique features. The vomeronasal organ in salamanders is found in a ventrolateral diverticulum of each main olfactory organ. P. cinereus has a more anteriorly placed vomeronasal organ within the diverticulum, and the posterior limit of each nasolabial groove is adjacent to the anterior limit of the vomeronasal organs. This suggests that the grooves deliver chemicals preferentially to the vomeronasal organs instead of to the main olfactory organs. In addition, the vomeronasal sensory epithelium is thickest anteriorly and is at its thinnest at about the level corresponding to the location of the vomeronasal organ in other salamander families. These adaptations suggest a specific mechanism of odorant delivery to the vomeronasal organ in plethodontid salamanders not found in other salamander families.