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Data organized chronologically. Sampling code: fs , specimens from field surveys conducted by us in gardens, nurseries and semi natural areas; sbp , specimens sent by people who knew our work through the information in social networks; bd , bibliographic data. To each new sequence a three digit numeric code was assigned. Loc codes are as described in Table 1. Data from a total of 13 domestic gardens, seven nurseries, two plantations all confined, humanized locations , and three semi natural areas humanized environments that are not confined and in direct contact with agricultural and forest areas have been analyzed Table 1.

In all three places recent habitat restoration activities have been performed, including the transplantation of autochthonous plant species from commercial nurseries. Locality codes correspond to those in Table 1. Amateur collaborators photographed the animals alive and fixed them in absolute ethanol. Specimens we collected were also photographed and external morphological characters recorded. Preserved specimens were examined under a stereo microscope and notes of their dimensions, appearance, color though this is affected by preservation , eyes, any stripes or pattern, the position of the pharyngeal aperture mouth and gonopore, if present, were taken.

Specimens with no visible gonopore were considered to be immature. It was possible to identify some specimens, even immature ones, to species level without further examination. For unrecognized specimens, or where identity was uncertain and required confirmation, a mature specimen evidenced by an open gonopore was selected and divided into various portions, being embedded in wax. The rest of the tissue is kept as voucher in the Genetics Department Universitat de Barcelona. PCR products of the 28S gene for some individuals, that yielded double bands in the direct sequences, were cloned using HTP TOPO TA Cloning Kit for Sequencing Invitrogen in order to be sure that only one type of sequence was recovered since the existence of a duplication of the ribosomal cluster is known in terrestrial planarians, Carranza et al.

The sequences of the clones showed that these bands corresponded to polymorphisms of one of the types. Seqman v. Mitochondrial coding DNA sequences were translated into aminoacids and aligned manually in Bioedit v. Hall, All sequences were unambiguously aligned. We estimated the DNA sequence evolution model that best fits the data for both molecules using jModelTest 2.

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Darriba et al. Ronquist et al. Bootstrap support BS values were obtained for ML trees from 10, replicates. In the BI analyses we ran four chains to allow heating and used default priors, three million generations were run using the Markov Chain Monte Carlo MCMC analysis in two independent runs. Sampling was every 1, generations. The stationarity and convergence of the runs were checked by plotting Log likelihood values vs. Using data describing the known distribution of C.

This could be a tool to help limit potential activities in order to avoid the introduced animals becoming invasive in the most likely areas for them to be successful. For the SDM, a total of Australian geographical coordinates of presence observations extracted from the literature, internet sources and personal communications L Winsor, were used for calibration of models training dataset. To avoid over-parameterization and loss of predictive power, we discarded the climatic variables that were highly correlated. To do this we extracted environmental information from 10, randomly generated points and determined the linear relationships among them using Spearman and Pearson correlations.

According to this analysis we used the 9 bioclimatic variables from the WorldClim database v. Those variables were: annual mean temperature; mean diurnal range; isothermality; maximum temperature of warmest month; minimum temperature of coldest month; precipitation of wettest month; precipitation seasonality; precipitation of wettest quarter; and precipitation of warmest quarter. The maximum entropy model, a presence-only algorithm that requires known species occurrence points and environmental variables Maxent v. We selected the software default values for the convergence threshold, regularization values, and features.

The maximum number of iterations was set to 1, and 1, bootstrap replicates were used. Once the models were trained, we projected the results using the IP climatic dataset, to study the possible expansion of C. Model performance was evaluated using the AUC test area under the receiver operating characteristic curve ROC and the binomial test of the omission-dependent threshold was calculated by Maxent.

Finally, binary maps of the outcome of the models were overlapped in the geographic information system, ArcMap v. Based on the external appearance of the flatworms we initially grouped the specimens into nine morphotypes. We classified four of them at the species level due to their characteristic shapes or other external features, and the other five at genus or tribe level. Bipalium kewense Fig.

A Dorsal view. B Ventral view of median part. C Dorsal view of posterior end. D Dorsal view of anterior end. Scale bar 5 mm. For Caenoplana bicolor Graff, there is no published description of a sexually mature specimen, hence the identification of the only specimen obtained, also an immature individual, relied exclusively on its external appearance Fig. This specimen is deposited in the tissue collection of the Genetics Department Universitat de Barcelona. Dorsal view with partial ventral view in the center.

The anterior end is not shown the specimen was damaged in this region. Among the specimens with an external morphology initially ascribable to the Caenoplana coerulea phenotype, we have found two morphotypes basing on their color pattern. Morphotype Ca1 Fig. Morphotype Ca2 Fig. C Lateral view of anterior end showing line of eyes.

B Ventral view of median part and dorsal view of anterior end showing line of eyes. Dolichoplana striata Fig. C Lateral view of anterior end showing the eye spot. Kontikia ventrolineata Dendy, Fig. B Ventral view of posterior end. We found one morphotype externally ascribable to the genus Rhynchodemus , but not to a known species Fig. Rhynchodemus morph Rs1 has a dark brown pigmented body with two black longitudinal stripes, and two large eyes situated a little distant from the anterior tip. A Dorsal view, scale bar 5 mm. B Lateral view of anterior region, scale bar 2.

A morphotype externally ascribable to the tribe Rhynchodemini was found in Benamargosa locality Loc-code G , but its morphological features did not allow assigning it to any genus. Rhynchodemini morph Ri1 presents a dark brown pigmented body with one dorsal black line no image available. Specimens of Obama sp. Externally, they are very similar to Obama sp. A Dorsal view of two specimens codes and from one Obama sp. A clade in the Geoplaninae Cox1 tree Fig. B Dorsal view of two specimens codes and from Obama sp.

B clade in the Geoplaninae Cox1 tree Fig. C Ventral view. D Lateral view of anterior end showing line of eyes. We inferred ML trees to check the diagnosis of the introduced specimens and to determine their level of relatedness to the ones from the original areas of distribution. For this reason, the datasets included, when possible, sequences belonging to morphologically diagnosed specimens from the original area of distribution of the putative introduced species obtained for this study or coming from GenBank; Table 2. We obtained 28S sequences for 15 individuals. One or two sequences from each morphotype were aligned together with 19 GenBank ingroup sequences and 3 outgroup sequences belonging to the Dugesia genus terrestrial planarians sister group; Carranza et al.

Cox1 sequences were obtained for all individuals included in the study Table 2. We inferred a ML tree with partitions from a concatenated dataset including 37 individuals for which both 28S and Cox1 sequences had been obtained Fig. The ML trees obtained from the Cox1 datasets are shown in Figs. Tree inferred from the concatenated dataset Cox1 and 28S genes. Three Dugesia species as outgroups. Tree inferred from the Cox1 gene. Three Microplana species as outgroups. One species of genres Arthurdendyus , Artioposthia , Australoplana and Caenopolana as outgroups.

For the concatenated dataset, the ML tree showed most introduced specimens constitute monophyletic groups together with representatives of their species coming from the original distribution area or other introduced localities. We have found introduced planarians in the IP for all non-autochthonous terrestrial planarians subfamilies; in the case of the Rhynchodeminae there are even representatives from two tribes Rhynchodemini and Caenoplanini.

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Within the Bipaliinae, Bipalium specimens found in the IP constitute a monophyletic group together with Bipalium sequences from other species, B. The genetic diversity among the four B.


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In the Geoplaninae clade Figs. In the Cox1 tree, specimens coming from the IP, United Kingdom both introduced and Brazil original area constitute a highly-supported monophyletic group. Within this group, the introduced individuals are divided in two quite differentiated clades Obama sp.

A and Obama sp. B in Fig. All the UK individuals fall within the clade Obama sp. Two Cratera species as outgroups. The Caenoplanini clade Figs. Even Caenoplana coerulea sequences, either coming from GenBank, or from the individuals sent by our collaborator in Australia, are found in very distinct genetic clades pointing to the existence of more than one species see Discussion. For this reason, we use the name Caenoplana coerulea s.

In the concatenated tree, the representative of Caenoplana morphotype Ca1 is closely related to Caenoplana coerulea s. The divergence among these three lineages can be appreciated when compared to the other subfamilies present in the tree.

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In the Cox1 tree Fig. This clade is sister to another group including C. The other two Caenoplana morphotype Ca1 individuals, coming from Townsville Australia , constitute a highly differentiated clade that also includes a GenBank sequence identified only to the genus level and one of the introduced individuals. Finally, there is a clade including only introduced animals, one of them identified as C. The genetic differentiation between the two lineages within this clade is nonetheless extremely high.

One Rhynchodemus species, one Platydemus species, and two Dolichoplana species as outgroups. In the Rhyncodemini clade Figs. Dolichoplana striata sequences form a monophyletic clade in the Cox1 tree, including three introduced animals in the IP and one coming from Brazil. The four K. The genus Rhynchodemus is represented by at least three species in the Cox1 tree.

It should be noted that the specific identification of all R. Rhynchodemus cf. In all the plant nurseries, only one species of terrestrial planarian was found Bipalium kewense, Rhynchodemini Ri1 or Obama sp. Obama sp. In the semi natural areas only the species K. The result of projecting models for the potential distribution of C. A composite map showing the potential distribution models for C. The color gradient indicates the predicted likelihood that the environmental conditions suitable for the species based on the MaxEnt average output.

Letters indicate localities where C. The results of the potential distribution of the species in the IP, based on data from its current distribution in their region of origin Australia , show that the species can find extremely suitable areas for its survival and expansion is the northern region, where the appropriate temperature and humidity conditions occur. External morphology Figs. However, the phylogenetic trees obtained and the analysis of the external appearance of the specimens indicate that probably at least five more species were present, including Rhynchodemini morph Ri1, Rhynchodemus morph Rs1, Obama sp.

There are no published Cox1 gene sequences for this species in Genbank, so those presented in this paper are the first available. Phylogenetic analysis of these sequences point to an introduction from the same lineage. Surprisingly, all sequences belonging to Kontikia ventrolineata coming from Spain, France and UK are situated within the Rhynchodemini clade with high support in both trees.

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This situation contradicts the taxonomy proposed by Sluys et al. The genetic differentiation observed within the group constituted by the genus Caenoplana , monophyletic in the trees, leads us to predict that it includes more than one species. In fact, C. According to Winsor, there are at least three species that are distinguishable morphologically; but there are probably more than three species in the area of origin. One of the problems with the group is that the type of the species is non-sexually mature. Hence, to clarify the situation and number of the species in this group, a broad sampling in its original area of distribution is required, followed by a thorough morphological and molecular study.

Nonetheless, for the purpose of the present paper, the evidence is clear that at least three different genetic lineages from Australia have been introduced in the IP, probably independently. In the case of Rhynchodemus Rs1, we cannot be sure if this is a distinct species or simply a differentiated lineage of R. The type locality of R. In our molecular analysis there was no separation of specimens according to their locality type natural or artificial. Two distinct clades of European Rhynchodemus were obtained Fig. In the case of Rhyncodemini Ri1, this species probably belongs to the genus Dolichoplana ; however, we were only able to obtain three specimens and none of them were sexually mature.

When specimens of Obama sp.


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Sampling performed in Brazil since then has found another species Obama sp. Molecular data show that it is closely related to the individuals found in Europe M Riutort, unpublished data, As in the previous case, a morphological and molecular study should be undertaken to clearly delimit and describe the new species.

The two clades found in our Cox1 tree Fig. Overall, we have shown that at least ten introductions have occurred in the IP. These introductions include species from all the non-European terrestrial planarian subfamilies from native localities as far as South America and Australia.

Since most of these species have previously been reported to have been introduced in other countries, the introductions into the IP have probably not been directly from the source countries, but were more likely to be indirect, following plant trade routes. In most cases, all the individuals from the same species found in the different localities are nearly identical, even when compared between Spain and the UK, which can be interpreted as the result of a single introduction or a single exportation from the place of origin.

In others, as in the case of Caenoplana , the observed diversity clearly indicates that the introductions were from different lineages within this group and is likely to be the consequence of more than one export from the native area. The feeding habits of the introduced species in the IP indicate that all of them feed on invertebrate soil fauna Table 3.

In plant nurseries and greenhouses microclimatic conditions are maintained artificially high humidity and stable temperature and are likely to favor the presence of stable populations of many species of terrestrial invertebrates. In nurseries we visited, especially under flowerpots, we have observed the presence of numerous specimens of snails, slugs, earthworms, millipedes, isopods, beetles and various groups of microarthropods, including springtails. Therefore, in this very suitable artificial microhabitat, there is likely to be a greater number of species of terrestrial flatworms as is the case of Bordils, Loc code V in Table 1 , where six species were detected in the same greenhouse.

The suitable conditions in the plant nurseries and garden centers may explain their introduction success.

In recent decades, the adoption of free market policies and trade agreements have reduced barriers to plant trade among different countries Dehnen-Schmutz et al. Depending on the intricate network of commercial interactions among European countries see Dehnen-Schmutz et al. Exotic species present in an area could be categorized as introduced detected in the area but with unknown status , adventives or not established they reproduce occasionally in the area not constituting stable populations , naturalized or established they form stable reproductive populations in the area and invasive established and well spread in the area Richardson et al.

The problem with this sort of assumption or calculation is that, in most cases, we simply have no knowledge of the unsuccessful introductions. In the case of terrestrial planarians, some species are very tolerant of habitat modification Cannon et al. In our case, most specimens occurred in confined areas gardens and nurseries. However, Rhynchodemus Rs1, C. In the particular case of C. The results show that the potential distribution of the species Fig.

The most suitable area is the northern IP.

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This is not surprising when we consider that in this northern region, the climatic conditions temperature and humidity are also more optimal for the presence of native land planarians Mateos et al. Thus, we show that by having suitable climatic databases, it is possible to model the potential distribution of introduced species, and thus predict their risk of becoming invasive.

If we add to this information the knowledge of some biological features of the terrestrial planarian species, such as their prey preferences, we may be able to make an even more precise image of the sites where it is more likely for the species to become invasive and thus concentrate prevention efforts in those areas. Our results show that C. This may be because it feeds on several groups of arthropods that are abundant in areas where this species has been detected isopods, beetles, diplopods. The three species Rhynchodemus Rs1, C. Another important question is: what are the negative effects of the spread of these species?

As predators of earthworms, planarians can cause soil drainage and fertility to be severely compromised. The ecological consequences of the presence of these predators depends on their propagation speed and efficiency, but could have significant effects on processes mediated by earthworms in both agroecosystems and forests Lilleskov et al. No reference has been made to the effect of these species on autochthonous populations of terrestrial planarians, probably because the knowledge of the autochthonous fauna is very scarce.

In the IP we have already performed some studies on the autochthonous terrestrial planarian fauna and found that it is very diverse, including at least 15 species, of which some contain a great deal of genetic diversity Mateos et al. The potential arrival of some of these introduced species in natural habitats, where the autochthonous ones are localized as predicted by the potential distribution studies , would have very negative consequences. Since exotic planarians are, in general, bigger in size, more voracious, have more aggressive behavior, and sometimes appear to have a generalist diet pers.

An important question raised by all these observations is whether governments in Europe should be asked to propose new, more restrictive rules on the trade of plants coming from outside, or alternatively, to establish better controls or protocols to avoid the introduction of unwanted organisms together with the plants. However, it is probably now too late to have an impact on the transport of species around the world.

Nonetheless, we are still in time to avoid invasions of terrestrial planarians. The restoration of degraded areas involves planting native plant species. These plants are available from nurseries and transported to the restoration areas accompanied by a certain amount of soil on the roots. If this land is not subject to any preventive treatment, it may be contaminated with organisms that are also introduced in the area that is being restored. Among these organisms may be unwanted species that, if given the right conditions, can become invasive. It is important to warn agencies conducting such restorations of these dangers and ask stakeholders to include in the protocols of landscape restoration the necessary steps to avoid these unwanted introductions.

Some simple, easy-to-perform sanitizing procedures, such as heating the soil EPPO, a ; EPPO, b ; SEERAD, ; Sugiura, before transplanting the nursery plants to the natural environment, may be sufficiently effective and reliable to ensure that there is no concomitant dispersal of flatworms. Such procedures, together with a periodic analysis of the introduced species present in garden centers and nurseries, and a study of the potential areas of flatworm distribution, would also help avoid the introduction of terrestrial planarians into areas where they are more likely to become invasive DEFRA, ; DOVE, Hannah Buckley, Dr.

Leigh Winsor, Dr. Jean-Lou Justine and an anonymous reviewer provided helpful comments that improved the manuscript. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. The following information was supplied regarding the deposition of DNA sequences:. See Table 2. Europe PMC requires Javascript to function effectively.

Recent Activity. The snippet could not be located in the article text. This may be because the snippet appears in a figure legend, contains special characters or spans different sections of the article. Published online Jun PMID: Corresponding author. Eduardo Mateos: ude. Received Apr 27; Accepted May This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed.

This article has been cited by other articles in PMC. Abstract Many tropical terrestrial planarians Platyhelminthes, Geoplanidae have been introduced around the globe. Introduction Most animal invasive species detected in Europe are terrestrial invertebrates Roques et al. Open in a separate window.

Figure 1. Distribution map of the terrestrial flatworms. Figure 2. M Species sectioned for internal anatomy study see Table 2. Sequenced specimens. Figure 3. Distribution of sampling localities of introduced terrestrial flatworms in the Iberian Peninsula. Morphological studies Preserved specimens were examined under a stereo microscope and notes of their dimensions, appearance, color though this is affected by preservation , eyes, any stripes or pattern, the position of the pharyngeal aperture mouth and gonopore, if present, were taken. Potential distribution modeling Using data describing the known distribution of C.

Results Morphological identification of the specimens Based on the external appearance of the flatworms we initially grouped the specimens into nine morphotypes. Figure 4. Bipalium kewense. Figure 5. Caenoplana bicolor. Figure 6.

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Caenoplana morph Ca1. Figure 7. The city of Berlin, population 3. Several municipalities large and small have experimented at different scales with municipal wireless services and have met with mixed success. But is municipal wireless Internet feasible for even smaller rural communities like Madison population 6, , Flandreau population 2, , and Montrose population ? It would seem that all communities could find benefits in open and pervasive Internet availability, but is there a lower bound to the size of community that can support its own citywide wireless network?

Municipal governments in cities the size of Madison often cannot afford to hire dedicated IT staff. Some smaller rural community governments hardly have the staff to maintain a municipal Internet presence, let alone build or administer a public wireless network. Yet in South Dakota, almost every one of these communities has a well-developed public computer network and Internet access, built through a cooperative local and state effort, in its school buildings and public libraries. The state supports a centralized administration system that coordinates Internet services for a couple hundred school districts spread over 76, square miles.

This paper investigates the possibility of expanding this system to provide municipal wireless Internet via WiMAX to every community in South Dakota and create a statewide wireless Internet hotspot. Today, being cut off from basic telecommunications services is a hardship almost as acute as these other deprivations, and may indeed reduce the chances of finding remedies to them.

Vos attempts to survey the number of municipalities with some sort of public wireless deployment. That survey finds jurisdictions with active region or citywide wireless programs, 74 jurisdictions with active city hotzones, and 55 jurisdictions with wireless networks strictly for public safety and municipal use. Vos also tentatively identifies jurisdictions with some sort of citywide or countywide wireless project in the works anywhere between request for proposals to deployment phase.

Cloud, Florida. While not a municipal system, the Mountain Area Information Network has operated since as an analogue to rural electric cooperatives and is expanding wireless broadband to rural areas, thus demonstrating the ability effective Internet service provision on a non-profit model. In an analysis of 25 different municipal wireless network projects, Mandviwalla et al. Aadikalam et al. To truly expand Internet access, communities deploying municipal wireless must also invest in education programs and public workstations. Mandviwalla et al. The public sector may also have a market interest in providing Internet service.

Telecom mergers are consolidating control of the Internet and other media in fewer corporate hands, raising concerns about price competition and network neutrality Noguchi, Some argue that a public-sector presence in the Internet service provider market adds competition and establishes a provider whose public obligation to treat all users equally would strengthen network neutrality Dingwall, The idea of expanding municipal wireless into a larger regional network has some conceptual of not practical precedent.

The city of Luxembourg established HotCity, a municipal Wi-Fi network that covers half the city and serves 12, registered users with Wi-Fi hotspots. The city plans to expand the network to serve 85, users with hotspots by the end of this year. Ultimately, says Mayor Paul Helminger, the city aims to expand HotCity to the entire country and its half million residents and conceivably to nearby French and Belgian cities Baritault, Nonetheless, Luxembourg appears to offer some useful lessons for establishing public wireless networks of municipal or regional scale.

Various technologies may be put to use to provide ubiquitous municipal Internet access.

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Wi-Fi networks can provide good coverage, but as Aadikalam et al. This shortcoming may limit use by emergency responders Aadikalam et al. Overall, the advantages of WiMAX over Wi-Fi appear to fit exactly the conditions necessary to bring municipal wireless to rural communities in South Dakota. A statewide collaborative network of community wireless networks certainly aims at ubiquitous and preferably mobile coverage.

Communities adopting this technology certainly hope for the problem of scalability as they tout the network to increase tourism, business, and even population. A community-wide network would need to support a wide range of government, commercial, and residential applications. Public agencies, businesses, and private citizens sharing a network would have reasonable concerns about security. WiMAX appears to better address those concerns. WiMAX is a line-of-sight technology well suited to the relatively flat urban and country landscapes of South Dakota.

The scarcity of major hills, river valleys, and forested areas will allow for signals from urban transmitters to reach numerous rural residents beyond municipal boundaries that, in many rural communities are well under a kilometer in radius. IEEE Sioux Valley obtains this reach with a single transmitter atop a twelve-story building, one of only four such structures in the city of comparable height the others include a grain elevator complex and two water towers. The relative lack of tall buildings in South Dakota communities means fewer urban dead spots; however, it also poses challenges for siting WiMAX transmitters for optimal reach.

Installing a WiMAX transmitter for municipal wireless at local schools, whose tallest structure may be a two- to three-story athletic complex, may require installing a radio tire on school grounds or arranging for extension of Internet backbone to the nearest grain elevator or water tower. One obstacle to establishing a statewide collaborative municipal wireless network through WiMAX may be frequency licensing.

A number of public school districts and universities in South Dakota hold educational broadband licenses for the 2. Unfortunately, these schools are clustered in the eastern third of the state, suggesting coverage for the less densely populated areas of central and western South Dakota may depend on communities or the state obtaining licenses for other frequencies. Yet it is worth noting that for those public license holders, putting their frequencies to public use in a WiMAX-based municipal wireless network would satisfy an FCC requirement that EBS license holders make some substantive use of their spectrum holdings for the public good by May 1, Teal, Another obstacle is access to the WiMAX signal.

Few if any home computers are WiMAX-compatible. A statewide deployment does gamble to some extent that the market will make available the necessary technology on the user end. There is the possibility, though, that a statewide deployment that follows a five-year timeframe like that of the turn-of-the-century project to wire the schools would coincide with market adoption of WiMAX receivers as standard wireless equipment on consumer electronics.

Rick Rotondo, chief marketing officer at Spectrum Bridge, says that EBS license holders might need to spend one million dollars to meet the minimum requirements for putting their licensed spectrum to use Teal, Of course, this estimate comes from a broker who has some motivation to encourage public and non-profit agencies to sell their licenses. Funding need not come strictly from the state budget; city and county governments, school districts, and even local economic development organizations stand to benefit and could justify diverting some portion of their IT spending or even designating a new budget item to support this project.

An extension of the statewide K network to support municipal wireless Internet would serve most immediately to expand the educational mission existing educational technology was implemented to carry out. The school-to-home connection was promoted in early discussions of the initiative to wire the schools as a way to improve student and parent engagement Janklow, Providing community-wide access to a unified network would facilitate the use of money-saving computing alternatives like netbooks and online software, which are already being explored by innovative school districts like Elkton and Madison Heidelberger, ; Clement, State support for a unified system of municipal wireless networks could jumpstart local efforts to establish online presences and functions for local governments.

Rural communities are interested in establishing an online presence that can serve public officials and citizens alike. However, without dedicated IT staff, they can hardly afford to implement major systems of their own. Municipal networks built and supported by a central state agency would free local agencies and citizens to work on ways to put that new infrastructure to good public use.

A statewide municipal wireless initiative has significant marketing value. Visitors-not just business professionals and academics, but increasingly regular working-class tourists-rely on e-mail and other Web-based communications and expect that connectivity when they are on the road for business or pleasure.