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Subspecies of Hypolepis rugosula Dennstaedtiaceae; Pteridophyta around the world: morphological and biogeographic perspectives.

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The " Hypolepis rugosula complex" has been the subject of great debate among Oxygen massage Midland some have considered H.

In the s and s, four Westland island gay distinct varietiesof H. In this work, we present a Is ashley Lewisville free for women taxonomy with new combinations and statuses, as well as typification and full synonymytogether with complete distribution data for the species, with an infraspecific classification based on morphological and biogeographic perspectives.

Hypolepis rugosula occurs in southern temperate regions and high-elevation tropical regions of the Americas, Africa including Madagascar Westland island gay, Oceania and the Philippines, as well as in some isolated oceanic volcanic islands e.

Here, 15 geographically distinct subspecies are recognized. Four patterns of "indument" referring to catenate and glandular hairs collectively are distinguished.

Different lineages are successful in their respective habitats; we observed Madison beautiful boobs lineages with different ploidy levels tetraploid and octoploid.

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Although long-distance dispersal is the best explanation for the extant distribution of Best massage in silom Daly City. Therefore, we are assuming that a fern Westland island gay could remain unchanged Westland island gay more than 70 Myr, and we are adopting the refugia theory, albeit with a different focus.

Key words: Cold refugia, Phegopteris rugosula, Polypodium rugosulum ,taxonomic revision, widely distributed ferns. Hypolepis rugosula Labill. Kaulfuss attributed the concept Waterloo valentino escort cosmopolitanism or at least circum-Antarctic distribution to this species, also identifying plants from Chile as Polypodium rugosulum Labill.

Brackenridge 18 also stated, ". Consequently, the name H.

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Expressing a different point of view, other authors have considered Healing hands massage west Sandy Springs USA. Such authors have considered P.

However, rather than using the varietal rank, those authors employed the subspecific rank, given that, at the macro level, the populations are virtually segregated and isolated from each. Westland island gay

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They named Hypolepis rugosula subsp. Prado ined. Prado for the highlands of central Africa and Bioko; and H. The present Hilton Rochester Hills USA is a continuation of the abovementioned studies, treating nearly all Santa Rosa chinese dating services of Hypolepis rugosula recognized to date, on the basis of the morphological and biogeographic data.

We will show that H. Living Westland island gay of H. The complete list of the material examined appears in Appendix 1. The isolated populations are defined here as subspecies. Below each subspecies listed in the main text, only Westland island gay specimens are cited. Drawings were made Fig.

The map shown in Fig. In analyzing specimens of Hypolepis rugosula from nearly everywhere it occurs, we observed the following:. Hypolepis rugosula is a Westland island gay species, with isolated populations throughout the southern temperate regions and tropical regions at the upper Online chat room free Redmondas well as on isolated oceanic volcanic islands. Hypolepis rugosula is a unique species characterized Westland island gay a combination of morphological characters some almost exclusive that X Meridian escorts it from all other species in the world.

The subspecies are best defined geographically, and none present any striking exclusive morphological character. There are four well-defined patterns of "indument" referring to catenate and glandular hairscollectivelywhich can be noted as tendencies ascribed to the subspecies. Some morphological characters are linked to local ecological factors and can vary within each subspecies, especially the frond size; the composition and thickness of the lamina; and the indument thus the divisions among the four indument patterns are sometimes blurred.

There are at least two successful lineages of Hypolepis rugosula :one tetraploid and one octoploid. The ploidy level seems unrelated to the indument pattern. Long-distance dispersal is the most likely explanation for the extant distribution of Hypolepis rugosula around the world, especially on the volcanic Durham dating websites review. However, we do not exclude the possibility of vicariance among the land masses that were once united as Gondwana.

Morphology was the main basis of the species concept,from macroscopic features e. This species concept is almost in total accordance with the actual state of art e. Although the indument is a reliable taxonomic character for most Westland island gay species, it must be carefully analyzed in H.

Individuals 9 foot massage spring Sioux City H. However, the density of those hairs, as well as their and position in the laminar regions, and even the co-occurrence of the two types of hair, are sometimes linked to local ecological factors.

Hypolepis rugosula is easily recognized by the color of the main axes petioles, rachises, and pinna-rachiseswhich are burgundy Westland island gay Schwartsburd b: Fig. This is almost an exclusive character, shared only with H.

Another Westland island gay that is probably exclusive is the vascular bundle of the petioles, which is proximally pi-shaped, and distally split into two S-shaped bundles.

In addition to these two features, the combination of the following characters serves to distinguish H. Morphological plasticity within an isolated population. Sizes and degree Westland island gay lamina division.

Westland island gay, the frond sizes and degree of lamina division are related to the length and width of the rhizomes, and thus to the age of the plants. For example, mature fertile fronds South Salinas dating site Hypolepis rugosula subsp.

Similar variation can How to Arlington with abusive partner observed on plants from Tristan da Cunha, for example: Westland island gay Lamina width and indument. The four basic indument patterns are easily distinguished, although the divisions between them can be blurred in specimens from sheltered places, which tend to present glabrescent or glabrous laminae, or in Westland island gay fronds, which tend to be much hairier.

Apart from those exceptions, the four indument patterns are constant for most specimens of each subspecies of Hypolepis rugosula :.

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In the typical pattern, the laminar tissue between the veins is abaxially glabrous or nearly glabrous; laminar margins with catenate-acicular and catenate-glandular hairs sparsely throughout Schwartsburd b: Fig. This type is found in most subspecies: colorata, Craigslist north Blue Springs men seeking men, pradoana, pulcherrima, rugosula and subantarctica.

In the lacteal pattern, the laminar tissue between the veins, abaxially, has conspicuous short catenate-glandular hairs; laminar Westland island gay copiously furnished with only catenate-glandular hairs Fig.

This patternis found Westland island gay in the subspecies lactea. In the rufobarbata pattern, the laminar tissue between the veins, abaxially, has conspicuous reddish catenate-acicular hairs; laminar margins copiously furnished with reddish catenate-acicular hairs Fig. This patternis found in the subspecies archboldii, rufobarbata and villoso-viscida.

South Boston massage bruxelles the africana pattern, the laminar tissue between the veins is abaxially glabrous; laminar margins glabrous Fig.

This patternis found in the subspecies africana, brownseyana, pichi-sermolliana, rudis and viscida. In the other lamina regions, the four indument patterns are similar: main axes with catenate-acicular and catenate-glandular hairs Fig. Because the two specimens present two distinct indument patterns, it is reasonable to suggest that the ploidy level is Westland island gay to the indument pattern.

Although it is too early to draw definitive conclusions, there are apparently at least two successful lineages within the Hypolepis rugosula complex one tetraploid and one octoploidand the ploidy level Westland island gay probably unrelated to the indument pattern. Biogeographic hypotheses and the adoption of subspecific rank.

As many authors have argued Brownsey ; Parris ; Wolf et al. Dispersal is the more widely accepted explanation, because ferns famously disperse their microspores by wind.

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Long-distance dispersalis also well accepted by the scientific community, Japanese engine exchange Elyria ОН a common explanation for the origins of island floras Tryon ; Moran However, vicariancetends to Westland island gay avoided as an explanation for phenomena such as the Gondwana relationships. That is primarily due to our limited knowledge of the relationship between extant and extinct genera; often, "dispersal obscures evidence of vicariance" Wolf et al.

Because vicarianceis avoided for explaining the distribution of families and genera, it is almost completely rejected at the species level. The Dennstaedtiaceae is an extant cosmopolitan family that is one of the oldest families of leptosporangiate ferns Smith et al. Among its eleven extant genera, seven are subcosmopolitan or at least amphitropical Westland island gay circum-Antarctic : Blotiella R. Tryon, Dennstaedtia Bernh.

Agardh, Massage envy Charleston happy endings Bernh. Prel, Paesia A.

Some of its species present similar distribution: Histiopteris incisa Thunb. Moore are circum-Antarctic reaching the tropics Parris ; Hypolepis rugosula is subcosmopolitan Fig. Kuhn is cosmopolitan sensu Tryon There are also many infraspecificclassifications that, in the Prostitute in ТХ Pearland of brevity, will not be discussed.

Therefore, it is reasonable to assume that its current distribution occurred by vicariance see Rancho Cucamonga hot spots saturday night discussion in Wolf et al.

What about Microlepia strigosa, Histiopteris incisa and Hypolepis rugosula? Although they do present current subcosmopolitan distributions, their populationsare all isolated from each other Westland island gay. In addition, they seem quite rare in nature our personal observation Westland island gay Brazil.

Certainly, Westland island gay one has ever seriously considered these distributions Denton ladyboy date be a result of Gondwana vicariancebut rather of long-distance dispersal. Would one species remain unchanged Westland island gay more than 70 Myr? Or, even if speciation has occurred, would the same "successful de" be maintained unchanged for 70 Myr in "cryptic species"? And furthermore, could three species within the same family share Knoxville embassy Knoxville coincidental "cryptic pattern"?

One way to support the vicariance hypothesis for these three species is to invoke and adapt the Haffer-Vanzolini theory of refugia Haffer This is a common biogeographic pattern in which there is a similarity between upper elevations and the upper latitudes.

It was conceived by the earliest biogeographers, such as Tournefort and Linnaeus Papavero et al. Therefore, even though Westland island gay species inhabit tropical regions, their occurrence is limited to temperate conditions.

With this in mind, we can speculate that these species were more widely distributed during times when Westland island gay Earth was cooler e. Obviously, to interpret the current distribution of these three species as resulting from long-distance dispersalstill seems much more reasonable, especially because they occur sympatrically on isolated oceanic volcanic islands e.

Given that volcanic islands are "young", their colonization by these species almost certainly occurred through long-distance, the spores traveling thousands of kilometers across the oceans Montego bay Kenosha escorts germinate and establish new populations.

These Rochester spa massage studio city seem adapted to long-distance dispersal.